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PHYTOPATHOLOGY
OFFICIAL ORGAN OF THE
AMERICAN PHYTOPATHOLOGICAL SOCIETY
W. A. Qrton
EDITORS
DoxALD Reddick G. p. Clinton
W. H. Barre
H. R. Fulton
Ernst Bessey
VVm. T. Horne
associate EDITORS
(\ W. Edgerton
H. T. Gussow
K. (\ Stakman
V. H. Stewakt
H. P. Harss
H. A Kdson
G. M. I^eed
J. 1^. HORKR
BUSINESS MANAGER
(-■. L. Shear
VOLUME VII
JANUARY-DECEMBER, 1017
With 11 Plates, and So Text Fujukks
• . •
> ■
• «
PriJI.lSIIKD FOIt TIIK SnciKTV
Williams & Wilkixs CnMr.v.w
I^ALTIMOHK, Ml).
254048
• •••
• • •
• •
••
• •
• •
• • • .• •
• • *• ••
•• •/ • • •
• • • •
• •
• •
CONTENTS OF VOLUME VII
No. 1. February
Jens Ludwig Jensen (1836-1904). F. K0lpin Ravn 1
The penetration of foreign substances introduced. into trees. W. H. Rankin. . 5
The generation of aldehydes by Fusarium cubense. Albert C. Lathrop 14 •
Does Cronartium ribicola winter on the current? W. A. McCubbin. . * 17
The injurious effects of tarvia fumes on vegetation. A. H. Chivers 32
Sour rot of lemon in California. Clayton O. Smith 37
A disease of pecan catkins. B. B. Higgins 42
Some new or little known hosts for wood-destroying fungi. Arthur S. Rhoads 46
Notes on Cronartium comptonisB III. Perley Spaulding 49
Further note on a parasitic saccharomycete of the tomato. Albert Schneider 52
Phytopathological Notes 54
Abstracts of paper presented at the Eighth Annual Meeting of the American
Phytopathological Society, New York City, December 26-30, 1916 58
Literature on American Plant Diseases 79
No. 2. April
The perfect stage of Glceosporium venetum. W. H. Burkholder 83
Puccinia subnitens and its aecial hosts. Ellsworth Bethel 92
Contributions to our knowledge of the white pine blister rust. W. A. Mc-
Cubbin 95
Species of Melampsora occurring upon Euphorbia in North America. E. B.
Mains 101
Recent cultures of forest tree rusts. James R. Weir and Ernest E. Hubert 106
A Rhizoctonia of the fig. J. Matz 110
Buckeye rot of tomato fruit. C. D. Sherbakoff 119
Noteworthy Porto Rican plant diseases. F. L. Stevens 130
Pycnial stages of important forest tree rusts. James R. Weir and Ernest E.
Hubert 135
Review 140
Phytopathological Notes 140
Report of the Eighth Annual Meeting of the American Phytopathological
Society. C. L. Shear 145
Report of the Berkeley Meeting of the Pacific Division of the American Phyto-
pathological Society. W. T. Horxe 150
Literature of American Plant Diseases 152
No. 3. June
Studies on Bacterium solanacearum. E. E. Stanford and F. A. Wolf 155
Sparassis radicata, an undcscribed fungus on the roots of conifers. James n.
Weir 166
CONTENW
Some ehanflet produced in strawberry fruits by Rhisopus nisrieans. Nbil £.
Stb%'bns asd Low A. Hawkins 17S
Witches-brooms on hickory trees. F. C. Stewabt ISo
A new leaf-spot disease of cherries. Bmwrr A. RuDOLm 1k8
Elister spot of apples and its relation to a disease of apple bark. Dban H.
Rose 198
The pathoKenic action of Rhisoctonia on potato. H. T. Guasow 209
Synthetic culture media for wood-destroying fungi. Ernest J. Pieper, C. J.
Ilmi-HRET AND S. F. ACRBB . 214
Phy topathological Notes 221
Literature on Plant Diseases . . 228
• No. 4. Ai'orHT
The cupraminonium washm. their preparation, biological properties, And
application. <). Ri'tler 2.'i5
IlacilluH fnorulanii N. Sp. A Imcterial organivm found associated with curly
top of the sugar brct. 1*. A. lio.vcqrET 2f>9
A now apparatus for aiicptic ultrafiltration. Hali-h K. Smith . 2tK)
Factors nffrcting the |>araititiiim of Cstilago Zco*. F. (i. Pieiieibel 294
Hclcrotium bataticola. the cuune of a fruit-rot of |M*pper8. William II. Martin .'tfK)
A Nc^trin pnrn^itic on Norway innplc. NIel. T. Cook 314
Phy topathological Notcii . 'M*\
Literature on Plant DiaoaNcs 32.3
No. 5. CKrroBER
Alternaria on Datura and potato. R. D. Randh 327
Suacrptibility of non-citrua plants to Hactcrium Citri. R. A. Jeiile XV^
SotiK' dii>r.'uM.*H of oconoiiiic planta in Porto Rico. L. K. Mileh . . . 3I.'>
Tm-o now fon*tit trre ruata from the northwcat. H. S. jArKrtoN. . .'i.V2
Rolatjiin of toiii|)eraturv to the gniwth and inf«*cting |>owcr of Fiutariurii Lini.
W. H. TiHiiALK 3.Vi
A HiiiipU- and efT<*rtiv<* method of pn»tocting citruH fruits against 8tom-<*n(i rot.
JiiiiN .M. HoiiKiiH A\i> F. S. Karlk 3r»l
Arthro|MMii« iind gafltcn)|>odM aa carri<*ni of ('ronartium ribicola in gn*€ii hc»uH<*H.
<;. FLirrn <;raVATT AND RrMII P. MAk.HIIALI 'M\S
Pri'iiiiiinary n*|Nirr on the vi*rtirnl di.*<tribuiifin of Fuaarium in aoil. .Minmk W.
TwLoK ;17I
Brii*f»T .\rtii'l<"*
Notr^ on WfMMl-ilifit roving fungi which grow im lM>th ronifi*rou?< un<i dcridii-
oii!t tr*N'Fi II. Jami:m H. Wcir 379
Thr "pr.'U iiii'thiHl iif :ip|ilying ronrrntratrd fonnaldi'liydt* solution in the
rontrol iif fint Htniit. K. J. IIahkkij. 3^1
RrvifHii 3h|
Ph\ lopnl holiiiiir.il \«itfi« .HsS
LitiTaturi' on IMinr Di^fuac^if . . ^V.*3
\t». (i. Dl.i'l.XlhKK
A twig%nd N'af diJ^iTi.-**- of K<-rri.i j:i|Ninir:t V. K. Stkw \nt 3*.#9
Thi; rromn r;ink«T ili-**.-!**' of r**-*' L M. .M^-^^^Ki 40**
Contents
An epiphytotic of cane disease in Porto Rico. John A. Stevenson 418
The effect of roentgen and ultraviolet rays upon fungi. H. L. Tbumbull and
J. W. HoTsoN 426
Sclerotinia Trifoliorum, the cause of stem rot of clovers and alfalfa. A. H. Gil-
bert AND C. W. Bexnett 432
The conduction of potassium cyanide in plants. John A. Elliott 443
Reviews 449
Phytopathological Notes 449
Literature on Plant Diseases 455
INDEX TO VOLUME VII
New scientific names &re printed in full-faced type
Abies, amabilis, 140; balsamea, 46, 314;
concolor, 140; grandis, 140, 379; lasio-
carpa, 379; magnifica, 140; nobilis, 140
Abronia, elliptica, 93; fragrans, 92; mi-
crantha, 93
Acalypha, bachmeroides, 160
Acer, negundo, 445; rubnim, 48; sac-
charin um, 48
Agree, S. F., see Piepbr, Ernest J.
Acrosporium, fasciculatum, 39
Actinomyces, scabies, 212
Adiantum, 35
iEcidium, Abroniae, 93; fumariacearum,
93
Ageratum, conyzoides, 160
Aegle, glutinosa, 59
Agriolimax, agrestis, 369
Agropyron, smithii, 73, 226
Alerodes, citri, 132
Alfalfa, leaf spot of, 70; lightning injury
to, 142; stem rot of, 432
Alsike, 70
Altemaria, 63; Citri var. Cerasi, 188;
crassa, 329; cucurbits, 196; Solani,
316, 327; on Datura, 327; on potato,
327
Amaranthus, blitoides, 92; retroflexus,
92
Ambrosia, artemisisfolia, 157; trifida,
445
Amelanchier, alnifolia, 109
American Ph ytopatho logical Soc'iety,
abstracts of papers presented at eighth
annual meeting, 145-149; Report of
the Pacific Division, 150-151.
Amsonia, 67; cliate, 69
Angular leaf spot, of cotton, 64; of cu-
cumbers, 61
Anthracnose, beah, 61; of camphor, 59;
cucumber, 62; of onion, 59; raspberry,
83
Aplanobacter, agropyri, 226
Apple, blister spot, 198; bitter rot of,
132; fruit-rot of, 59; Hypochnus
ochroleucus on, 130; root-rot of, 77,
223; rot of, 76; rough-bark or scurfy-
bark canker of, 202; scab, 76, 221
Arachis, hypogsa, 156, 160
Arbutus, menziesii, 380
Armadillidium, vulgare, 369
Armillaria, mellea, 175
Arthropods, carriers of Cronartium ribi-
cola in greenhouses, 368
Aschersonia, on Guava, 132
Aspidium, 35
Aster, 67
Atriplex, canesccns, 93; confertifolia,
93; hastata, 92
Auerswaldia, palmicola, 131
Avocado, Mycosphserella persese on, 350;
Phyllachora gratissima on, 350
Azalea, viscosa, 32; effect of tar smoke
on, 32
Bacillus, amylovorous, 75, 200; calif orni-
ensis, 285; coli, 200; coli communis, 14;
mesentericus aureus, 285; morulans,
269
Bacterium, Citri, 58, 339; Dianthi, 284;
herbicola aureum, 285; Malvacearum,
64; mesentericus, 159; Phaseoli, 61;
solanaceanun, 155; viridilividum, 392
Bailey, Ernest, see Jones, L. R.
Banana, disease of Cuban, 14
Barley, blight, 69
Bean, anthracnose, 61; blight, 61, 65;
Cercospora canesccns on, 349; Di-
morium grammodes on, 131 ; Isariopsis
on, 345; loaf blotch of, 345; loaf spot of,
345; mosaic, 60, 61; root rot, 61
Begonia, effect of tar smoke on, 35
Bennett, C. W., see Gilbert, A. H.
u
Index
Bethel. Kllawortii, Pucciniasubiutens
and itjt a<>cial luMta, 92--4M
B«tula. lutca, 47. 48; occidcntalis, 379
BidcDfl. hipinnata, 163
Birch, vHlow. 4K
BiABY. ( I. H.. The nhort -cycled rromyccs
of North America (abstract), 74
Bitter rot, of apple. 132
Bjerkandera. aduMta, 47
Black heart, of orange, 190
Black H|Mit. of |M>pf>er. (>3
Blakk. M. a.. Cook, Mkl T., and
H(*HWARZE. (\ A., Studies on peach
yellows and little |>each (abstract), 76
Blatta. orientalis. 370
Blight, on barley. 69; on liean, 61, 65;
on rhcHtnut. 313; on eggplant, 60, 78;
on jM'arj*. 7.5; on |M)tato. 374
BlistfT }«|M»t. of HftplcM. 19S
Blurne:t. ImlHainifrra. HiO
Boletus. rfir(»in.'i|N>H. 74; frost ii. 74; gra-
cilij*. 71; ind('t«t^UM. 74; purpureus, 74;
s(>eriiif«UM, 74
B()N'(iri:T. V. A.. Baeilluj* nionilans \.
Hp.. 2iM> 2s«»
BotrvtiH. rin«T«':i. '.i*M
Brawiira. oliTacru var. aeephnla. 141
Bread fruit, runt of. 131
Br<m K. \\ . .*^.. and HiTH. W. A.. Con-
trol of ;ipplr rtrub by bleaching [m>w-
der :ib-tr.irt •. 7»i
Broniuh. h<Mikcrianu.*<, 14.3; marginatur*.
60; Mtrhi'uj*!?*. ♦'»•.*
BiMMiks. Cii\HLKs. and ('o<»LKY, J. S..
Jf>nathnii Mfwit •jtlM*tnict >. 7i): TfUi-
fienttun* n*l:ition.*« of apple rot fungi
(abstract . 7»»
Brom.'iili.'t. drtiiiNftn. 161
Bhows. Xh.i.ik a., a b.'ict<TijiI Hti'in and
leaf iliM'aj'r of li'ttuci* al»Mtra<*t ■, <».'{
Bt'KKii<»i.i»» i(. W. I!.. Ii«'an «lis<'aj««*M in
Nm ^ ork St«t«* in n»l»i abstract •, 61;
TIm» jK-rfrrt f*tag»* of < tlii-oM{M>riuni
vem-tuin. M •»!
Bl'Tl.l.H ( K . rh«' ('(ipraiitiiioiiiuni u ;u*h«*M.
Zl'f Jt;s
Buttf-rnut. 17
Byakn. L I* . 'rvl<*firliu«» tntiri on \%hfat .
•Vi; n ni'liiatodi- di*i'a-«* of t\\r da**h«>«'li
and it» miitrol \t\ hot uatt-r tr«-at-
nirnt abftrart . «»»i
Cabbage, wilt, 375
Calendula, officinalis, 32
Calonyction, 67
Camellia, sooty mold on, 133
Camfmnula, 68
Camphor, anthracnose of, 59; limb can-
ker of, 59
Canfiytuft. effect of tar smoke on. 32
Canker, of camphor trees, 59; citrus, 58;
Illinois, 205; rough bark or scurfy
bark, '202
Canna, rust on, 132
Capno<lium, brasilicnsc, 134;coflreie, 134;
footie, 134; mangiferum, 134; stella-
tmn, 134
Capsella, burMa-pastoris, 92
Ca|)«icum, annuum, 160, 312
(^arica, papaya, 349
Carpinus. 314
Carrot, lightning injury, 142
C'akhn'kk, Ktbanks, Do the bacteria of
angular leaf spot of cucumber over-
win t<'r on the seed? (abstract), 61
Carya, illinoenHis, 42; ovata, 185
('aasava, CercoH|)ora hcnningsii on, 349
(*astanea, dentata, 47
Cai4t<»lleja, angustifolia, 106; miniata,
C<*h'ry, bartrrial heart wilt, 64; crown
rot. t»l; croun n)t wilt, 64
Ct'phuh'ururt, virrHceni4, 1.32
Crrcospora. Apii, tV5; cancHcenn, .349;
rotTra-. .'ill*; craasa. .'i.*i5; cruenta, .'M6;
Datura*. .Ti.'); hcnningrtii, 319; Hibisci,
.'<r.»; IbiridiH, ,'i.37; Lunaria', .'i.37; \ico-
tiaiw. .34s ; Solaninigri, .TJ7
CfTcal pathologists, field conference of,
4,7.3 4.'>4
Charcoal rot. of Hweet |K)tatoe«, 312
ClK'noiNNliuin. alburn, 92; glaucuni, 92;
lanceoliituni, \f2', pagonuni, 92
Cherry, h'af H|>ot, 7.*), ISS; witches-
br<N>rn.*4 on. 1V»
ChcMtnut. Iilmht. 313
Ciii\Mt>«. A. II.. The injurious efTectM of
T.-ifMi furnr.M on vegetation. 32-»'ki.
Chr\>Miri\ \.i. Abietin, 78, XVJ; I*iceie,
.'CiJ. wiirii. ii.K{ .
( 'hr>-«»pM.M. 67; niariana, 6S
CitruM, trifoliata. .VJ, .*i.'J9; canker, 58
Index
ui
Cladosporium, Citri, 60; cucumerinum,
62
Clavaria, amethystina, 171; aurea, 171;
formosa, 171
Cleome, serrulata, 93
Clover, leaf spot of, 70; Nigredo fallens
on, 70; rust, 75; stem rot of, 432
Cnicus, americanus, 44
Coccomyces, hiemalis, 75, 404; KerrlsB,
405
COERPER, FLOR*feNCE M., SCe JOHNSON,
A. G.
Coffea, arabica, 115; liberica, 115
Coffee, Cercospora coffeae on, 349; dis-
eases of, 130
Coleosporium, Apocynaceum, 69; deli-
cat ulum, 68, 225; Elephantopodis, 66;
Helianthi, 67: inconspicuum, 68; Ipo-
mseae, 67, 132; Laciniarise, 68; Solida-
ginis, 20, 68, 225; Terebinthinaceae, 67;
Vemoniae, 67
Colletotrichum, cereale, 450; circinans,
59; falcatum, 423; fructum, 59; solani-
colum, 226
CoLLEY, Reginald H., Mycellinm of the
white pine blister rust (abstract), 77;
Pycnial scars, an important diagnostic
character for the white pine blister
rust (abstract), 77
Collins, J. Franklin, On using an
ether freezing microtome in warm and
damp weather, 222
Colocasia, esculenta, 66
Commelina, nudiflora, 35
Comptonia, asplenifolia, 49, 108
Conifers, Sparassis radicata on, 166
Convolvulus, 67
Cook, Mel. T., see Blake, M. A.; and
ScHWARZE, C. A., Apple scab on the
twigs, 221; A Nectria parasitic on
Norway maple, 313-314.
Cooley, G. S., see Brooks, Charles
Coreopsis, 67; dclphinifolia, 69; lanceo-
lata, 69; major, 69; major semeri, 69;
major rigida, 69; tripteris, 69; verti-
cillata, 69
Coriolellus, sepium, 47
Coriolus, abietinus, 46; nigromargina-
tus, 46; prolificans, 46; versicolor, 46
Corn, Phyllachora graminis on, 55, 131;
smut, 73
Corticium, javanicum, 115; Isetum, 115;
ochralcucum, 113; salmonicolor, 111;
vagum. 111, 209
Cotton, angular leaf spot, 64; lightning
injury, 140; rust on, 133
Corylus, avellana, 52
Cratsegus, douglassi, 109, 223
Cronartium, cerebrum, 135, 450; coleo-
sporoides, 106, 135; Coiflandrae, 135;
Comptoniae, 17, 49, 106, 135; Quer-
cuum, 20; ribicola, 17, 58, 77, 95, 135,
225, 368. 391, 449
Croton, glandulosus var. septentrional is,
161
Crown canker, of rose, 408
Crown rot wilt, of celery, 64
Crown rot, of celery, 64
Cucumber, angular leaf spot of, 61;
anthracnose, 62; mosaic on, 61; new
leaf spot, 62, scab on, 62
Cucurbita, maxima, 32; pepo, 32
Cuprammonium washes, 235
Curly dwarf, of potato, 71
Curly top, of sugar beet, 269
Currant, Cronartium ribicola on, 17, 58;
effect of tar<smoke on, 32; rust on, 17,
225
Cycas, revoluta, 44
Cylindrocladium, 410; scoparium, 409
Cylindrosporium, onKerria japonica, 399
Cystospora, batata, 74; Sacchari, 424
Daedalea, confragosa, 379; heteromor-
pha, 380; unicolor, 379
Dalbby, Nora E., Corn disease caused
by Phyllachora graminis, 55
Dale, E. E., see Melchbrs, L. E.
Damping-off, of tomatoes. 319
Dasheen, nematode disetis? of, 6(5
Datura, cornucopia, 160; fastuos'i, 160.
327; ferox, 327; incrnus, 327; licvis,
327; metelloides, 160; quercifolia. 327;
stramonium, 327; stramonium gi>?an-
teum, 327; stramonium inermis, 327;
tatula, 65. 163. 327; tatula inermis. 327
Davis, W. H., and Johnson, A. G., The
aecial stage of the red clover rust
(abstract), 75
^MAMhl
IV
Index
Dianthuii. incamata, 289
DiEHL, William, see Melhur, I. K.
Dimerium, fcrammodes, 131
Diplocladitiin, 410; cylindroflponun, 410
Diplodia, cacaoicola, 423; Camphonp,
60; rot of citrim fruits, 301
Distichlifl, spicata, 02
Dothichiia, populoa. 4^
DiRRELL. W. L., Notos on curly dwarf
nymptomson Irish potatoes (abstract),
71
Karl, F. S.. seo Kcmikrs, John M.
Karly bliKht, on Datura, 327; on potato,
327
I'}chino<iontium. tinctoriurn. 170
Kclipta, alba, 1.57
KixiEKToN, (\ W., A (rnomonia on ogj?-
plant (abstract). 78
Kdhon. H. a., nn<i S<-iirki.\kr, Ohwald,
A malnutrition diwiiMo of the Irish
|>otato and its control (abstract), 70
I*>lwomi. attacking the dasheen. 00
Kionilant. ( -olletotrichum solanicoluin
on. 2'2i)', (»nomonia on. 7K; Lifchtnin^
injury to. 1 10; stem blight of, 00. 78;
wilt on. 227
Klephantopus, tomentosus, 07
KlfvinKta. mcKaloma. 47
Ki.LioTT. John a., A new parasitic slime
mold suitable for class work (alwitract ),
74: Thi> conduction of |H)tassium cya-
nidi* in plants. 44.'i-448.
Klymus. americanus, 143; canadeiutis,
73; condensatiis, 73; rIaucus. 09, 73;
macounii, 7Ii; nibustus, 0!)
Kndothia. |>arasitica. 313
F'riKcnm. ranad«*nsis, 10.3
Krvsiniiiin. as|N*rum, 92
Kupatoriijin. OS
Kuphorbia. lOl ; commutata. 101; rypa-
rissia}*. 101: rxif^ua. lOl; falcata. UT2;
fCcrardiaria. lOTi: mutatis, lOJi: |M*pliiH.
lot: riibuHta. 101
Kuti»tti\. t«'ni'lla. -7:{
Kuthaniia. OS; r:ir<»litiiari.'t, OS; Icpto-
ri'phala, *kS
K\«»:i5»ru«i, <'«Ta»i. Isji
FAfLWRTTIK, It. <\, l)l«».»i«iiiiiiul|on «if
Bacterium Malvacearum (abstract),
04
Fern, effect of tar smoke on, .34
Ficus, 133; carica, 110
Fields, W. S., Method for photograph-
ing plate cultures, 388
Fig, Rhisoctonia of, 110
Flax, 359; wilt, 375
Fomea, annoeus, 175, 210; applanatus,
217; pinicola, 210, 420
Forest trees, rust on. 100, 135
Formaldehyde solution, for control of
oat smut, 381
FraRaria, .32, 178
Fromme, F. D., Tylenchus tritici on
wheat in Virginia, 452
Fromme, F. D., and Thomas, H. E., A
Xylaria root-rot of the apple (ab-
stract), 77
Fruit rot, of pepj>ers, 308; of tomato. 00
Fi'LTON, If. H., Manual of Fruit Dis-
eases, Hehlkr. Lex U. and Whetzel,
Hkrhert HirK, (Review), :W0-388
Fusarium, conglutinans, 358; cubense,
14; distribution in soil, .374; Lini, .3.50;
martii, 01 ; oxys|K)rum, .375; radicicola,
.375; roseuin, 384; Solani, .375; subula-
tum, .385
(Ialloww. H. T., Newton. H. Pierce.
143
(ttiniMlerma. Tsug:e. 47
(lARDNKR. M. \V.. I)iss4'mination of the
organism of cucuml)er anthracnose
tal>stract), 02
(Iasten>fK»ds, carriers of (Vonartium
ribicola in gn*enhous«>s. .'(08
(teranium, effect of tar smoke on, .35
(liblMTella. saubinetti. .'i8.">
(iiLHKRT. W. \V.. Virulenrr of different
strains of C-ladoMporium rucumerinum
(alwtrart :, 02
(tiLiiKRT. A. H. and Hknnktt. C W.,
.*>el«' rot i Ilia Trifolioruiii. the caiisi* of
hti*ni rnt of eloviT" and :ilfalfa, 4.32-
112
<;l\|)\%i\. v. K. aiicl HKDhh'K, Donald,
Siilfuritiic <'«>ne«»rd grain's t<i pn'vent
|N)udrry inildi'U 'abs^trart •, ♦!♦»
(il<ro|M)rus. coriehoiih'S. IS
Index
V
Glopophyllum, hirsutum, 48; trabeum, 47
GlcDosporium, Camphorse, 59; Mangi-
ferffi, 132; venetum, 83
Glomerclla, 85
Gnomonia, on eggplant, 78
Gondule, rust of, 132
Grapefruit, canker on, 59 i
Grape, powdery mildew, 66; rust on, 132
Graphiola, Phoenicis, 131
Grasses, rust on, 73; Scolecotrichum
graminis on, 69
Gravatt, G. Flippo and Marshall,
Rush P., Arthropods and gasteropods
as carriers of Cronartium ribicola in
greenhouses, 368-373
Guava, diseases of, 131
Gt'ssow, H. T., The pathogenic action of
Rhizoctonia on potato, 209-213; The
occurrence of Colletotrichum cereale,
Dothichiza populea and Leptosphseria
Xapi in Canada, 450; A thumb clip for
use with magnifiers, 451-452
Gymnosporangium, nelsoni, 109; tubula-
tum, 109
Hapalopilus, gilvus, 47
Haskell, R. J., The spray method of
applying concentrated formaldehyde
solution in the control of oat smut,
381-383
Hawkins, Lon A., see Stevens, Neil E.
Hedgcock, Geo. G., Edible and poison-
ous mushrooms. W. A. "Murrill
(review), 140; and Long, W. H., The
aecial stage of Coleosporium Ele-
phantopodis (abstract), 66; and Hunt,
N. Rex, The Peridermium belonging
to Coleosporium IpomoDse (abstract),
67; X Peridermium belonging to Co-
leosporium Terebinthinaceaj (ab-
stract), 67; An alternate form for
Coleosporium Helianthi (abstract),
67; Some new hosts for Coleosporium
Solidaginis (abstract), 68; Some new
hosts for Coleosporium inconspicuum
(abstract), 68; Notes on Razoumofskya
campylopoda, 315
Helianthus, 67; annuus, 321, 445; aus-
tralis, 68; decapetalus, 68; divarica-
tus, 68; eggertii, 68; giganteus, 68;
. grosse-serratus, 68; hirsutus, 68; mi-
crocephalus, 68; saxicola, 68
Helminthosporium, mayaguezense, 351
Hemileia, vastatrix, 130
Hemlock, sap-rot in, 47
Heterodera, radicicola, 66
Hickory, 44; witches-brooms on, 185
HiGGiNS, B. B., A disease of pecan cat-
kins, 42-45
Honeysuckle, effect of tar smoke on, 32
Hordeum, jubatum, 69, 73, 143; nodo-
sum, 69
HoTsoN, J. W., see Trumbull, H. L.
Ho WITT, J. E., Phytopthora infestans
causing damping-off of tomatoes, 319
Hubert, Ernest E., see Weir, James R.
Humphrey, C. J., see Pieper, Ernest J.
Humphrey, H. B., Puccinia glumarum,
142
HUNGERPORD, C. W., SCe JOHNSON, A.
G. ; Puccinia graminis on wheat ker-
nels and its relation to subsequent
infection (abstract), 73
Hunt, N. Rex, see Hedgcock, Geo. G.
Hypochnus, ochroleucus, 130
Iberis, amara, 32; umbellata, 337
Impatiens, balsamina, 161 ; sultani, 163
Indigofera, arreeta, 160
Inga, laurina, 132
Inonotus, dryophilus, 48
Ipomcea, batata, 312; lacunosa, 67; pan-
durata, 67; triloba, 67
Irpex, lacteus, 379
Isariopsis, on bean, 345; griseola, 345;
leaf-spot, 345
Ischnoderma, fulignosum, 47
Jackson, H. S., A species of Chrysomyxa
new to North America (abstract), 78;
Two new forest tree rusts from the
northwest, 352-355
Jagger, Ivan C, Two transmissible
mosaic diseases of cucumbers (ab-
stract), 61
Jehle, R. a.. Citrus canker investiga-
tions at the Florida Tropical Lab-
oratory (abstract), 58; Susceptibility
of non-citrous plants to Bacterium
Citri, 339-344
VI
Index
Jbn'SEN, Je\h LrDWiu. Biography of. 1
Jimson weed, leaf-Hpot of, 327
JoiiN'fuiN, A. 13.. flee TArBENiUNR. J. J.
JoiiXfM>N. A. <t. and ("obkpck, Flor-
ENCK M., A bnctorial blight of soy
bean (abntract ), (&5; and HrNOERPORO,
C W., Scolecotrichum graminis on
timothy, orchard graas. and other
gramea (abstract). 69; see Joneh. L.
R.; see Da vim. \V. H.
Jonathan spot. 76
JoNErt. Fred Reuel. The Pseudopexiza
leaf spot diaeaiieH of alfalfa and red
clover (abstract). 70
J^.VEM. L. R.. John HON. A. (r. and
Reddy. C. S.. Bacteria of barley
blight «e4»d-bome (abittrnct). 60; and
Bailkv. Krnrht. FrcMt necronis of
pf>tato tiib«*ni (Hlmtrnrt). 71; Light-
ning injury to Knl«>. 14(^142
Juglanit. cinen*.!. 47
Juni|M»ruH. commiinii*. 1(K): Hco|>uloruin.
10!) : virginianii, 46
Kale, lightning injury to. 140
Keitt, <J. W.. S<M*<infl pro^n*HH rf|M)rt on
invcMtigtitiunH of Imf H|Nit f»f chorrieH
an<l pluriiM in WiHcnuHiii uilMtnict •. 7.*)
Kerri:i. j:i|Hinif':i. •'{{>!<
Kochi:i. Mcopiiria. *.^2
KRftt'T. \V. S., Ii:i<*ti*ri:il (IiK«*tiM«>H i»f
ri'Irry .ibsfnirf . t*\: \Viiit4*nng of
S'pforiri |M'f ro«*«'liti.i v.'ir. Apii i:ih-
Htr 11*1 '. ti.'i
Ku**hiii-ol:i. I'iri. I.'U; < lof*-*ypii, l.'kJ
Kuiii<|U:it. r:ink('r on, .V»
I^ariiii.'iri.'i. 67; rjirlri. 6h: «*lt*g:in!4. i\S;
rloiigat.'i. t»H: pHUnflorn. 6S; MC'irioHji.
6s : MTi»tiii:i. i'ts
I^arix. i-iin»iM»:i. Uis. .'{.V»: liiririiia. |0*»;
f »rr K It'll trtliM. UWt. 17»i. .'il.'i. .'l.V»
L%Tiiii<>i*. Ki.iiiJtT (V. Th*- gfiuTHtion of
aldfhv<l«'«< by Ku-.'irtuiii nikN^rifM'. 14-16
I.^f MiKht. of tig. Il'i
Ix*.tf bl<t*rh. of iMMfi. .{|-'i
Id'tii'TMnt. of U I II (it wht'.'lt. 2JI
lii'af ••iNif. nlf;ilf:i. 7M; U-an. Hi'r, rhiTry.
7.'i. Iss. «'liivir. 70: curiiiiibtT. «ij.
Jtiii?«i>ii ui'ftl. .(.'7: phifii. 7.'i: |Mit.ito.
•i.'7, toin:tti>. A27
Lemon, brown rot. 37; canker on. 59;
scab, 00; sour rot. 37 ; stem-end rot. 363
Lentinus, 171
Leniitcs. heteromorpha, 380;scpiaria, 217
liCpidium, densiflorum, 02; medium, 92
liCptoHphvria, Napi, 450; Sacchari. 131,
424
Lettuce, rot, 392; Sclerotinia libertiana
on, 60; stem and leaf disease of. 63
Levin. Kxra, Control of lettuce rot,
392-393
Lightning injury, alfalfa, 142; ca^-mts,
142; cotton. 140; eggplant. 142; kale,
141; potatoes, 140: radishes. 142;
sugar IxH't. 141; to sugar cane, 317;
tomato. 142: turnips. 142
Lima Ijoan. mtNtuic on, <K)
Lime, canker on. 59
Literatum on plant diHeases. 79-SI ; 152-
154; •22S-2:M; 3*22-326: ;««-.««; 455-4450
LoNti, W. If., Hi»e Hkimmim'K, (Jko. (1.
Ij4mic<*ra. tnrtarica. 32
Lunar ia, biennis. .'i.'i7
Lyco|N*rHicum, cerasifonne. 161 ; esculen-
turn. 52, 155; pyrifonne. 161
MarnH|M)rium. cookei. 32S; I>atunr. :i3l;
S«>lniii. 32S
Mains. K. li.. .SjHTii'H of MeluinfMora oc-
curring U|Mm Kuplif»rbia in North-
America. 101-105
Maij^mA, J. It.. w»e Stkvkns. F. L.
Muiidarin. ranker on, 59
Mango. Meliola on. 132
Manihot, IXi; utillisma. 349
Maple. 4-H: Nectria paraHitic on Nor-
wav, 313
Mar:ii«inius. S.icchari. 42 1
Marigold. efTi*ct f»f tar sinok«* on. •i2
Makhiiall. Rihh v.. »<•«' (;i{\v\TT. <l.
Flippo
Maktix. Wii.i.i \\i If.. \ S<*hToriiini di?*-
v:\M* of |H'p|HT«< ;ib'»tr;irt i, IV-I ; SrK*n»-
tiuin li:il:itirol:i. :{i)s 'M'2
M:irtyiii;t. proliowriilfa. pil
M\«*'»»:v. L. M.. The crown c:ink«T •li?«-
f:i?*i' «if row. Mis 417
M%T/. .r. A Uhiz<M*toni:i of thr fig.
110 lis
Index
vu
McClintock, J. a., Economic hosts of
Sclerotinia libertiana in tidewater
Virginia (abstract), 60; Lima bean
mosaic (abstract), 60; Will Spongo-
spora subterranea prove serious in
Virginia? (abstract), 72
Myrica, calif ornica, 49; carolinensis, 49,
108; cerifera, 49; gale, 49; inodora, 50;
pumila, 50
McCuBBiN, W. A., Does Cronartium
ribicola winter on the currant? 17-31 ;
Contributions to our knowledge of the
white pine blister rust, 95-l(X); and
Posey, G. G., Development of blister
rust aecia on white pines after they had
been cut down, 391-392
Melampsora, 101; Euphorbiae, 101; Eu-
phorbise-dulcis, 101; Euphorbiae-Eng-
leri, 101 ; EuphorbisB-Gerardianae, 101 ;
Gehnii, 101; Helioscopise, 101; Me-
dusae, 108; monticola, 103
Melanconium, Sacchari, 423
Melchers, L. E., and Dale, E. E.,
Black spot of pepper (abstract), 63;
Puccinia triticina Erikss. Leaf-rust
of winter wheat causes damage in
Kansas, 224; see Potter, Alden A.
Melh^s, I. E., Notes on mosaic symp-
toms of Irish potatoes (abstract), 71;
and Diehl, William, The develop-
ment of the aecial stage of Nigredo on
red clover (abstract), 70
Melia, 133
Meliola, Camellaj, 134; Citri, 134; fur-
cata, 131; Mangiferae, 132; Psidii, 131
Methods, for photographing plate cul-
tures, 388; for the differentiation of
pathogenic fungi in the tissues of the
host, 389
Microtome, ether freezing, 222
Microstroma, 132; album, on oak, 42;
Juglandis,- 185; Juglandis var. robus-
tum, 45
Miles, L. E., Some diseases of eco-
nomic plants in Porto Rico, 34:V-351
Mistletoe, 315
Monolepis, nuttalliana, 92
Mosaic, on cucumbers, 01; on liiiui bean,
60; on pea bean, 61; on potatoes, 71,
72; on muskmclons, 01
Mucuna, 160
Murphy, Paul A., Seed potato certifi-
cation in Nova Scotia (abstract), 72;
The economic importance of mosaic of
potato (abstract), 72
Murraya, exotica, 339
Mushrooms, edible and poisonous, 140 •
Muskmelons, mosaic on, 61
Mycorhiza, 74
Mycosphaerella, Perseae, 350
Myriogenospora, 132
Nasturtium, wilt of, 160
Naumov, N. a.. Intoxicating bread
(review), 384-386
Necrosis, of potatoes, 71
Nectria, <;occinea, 314
Nematode, disease of the dasheen, 66
Nematospora, Coryli, 52; Lycopersici, 52
Nicotiana, 160; tabacum, 155
Nigredo, fallens, 70
Norton, J. B. S., Host limitations of
Septoria Lycopersici (abstract), 65
Nummularia, discreta, 205
Oak, 42, 74
Oat, smut, 381
Oberly, Eunice R., (compiler), and
Smith, Florence P., Literature on
American plant diseases, 79-81; 152-
154; 228-234; 322-326; 393-398; 455-458
O'Gara, p. J., Notes on the distribution
of the bacterial disease of western
wheat-grass, 225; The occurrence of
Colletotrichum solanicolum O'Gara on
eggplant, 226
Oidium, citri-aurantii, 39; fasciculata,
39; tigitaninum, 39
Oospora, citri-aurantii, 39; fasciculata,
39; lactis, 40
Okra, Cerocospora Ilibisci on, 349
Onion, anthracnose, of, 59; pink rot of,
59
Oosporoidea, citri-aurantii, 39
Orange, black heart, 190; canker on, 59;
sooty mold on, 13.3
Osner, CJko. a., Preliminary notes on a
• new leaf spot of cucumbers (abstract),
62
VUl
Index
Palm, diseaBes of. 131
PantomuruB, fulleri. 369
Papaw, Pucciniopsifl Caries* on, 349
Parsley, Hclerotinia libertiana on, 60
Parthcnium, intern folium, 67
Paspalum, 132; conjugatum. 351
Peach, brown rot, 179; little, 76; yel-
lowfi, 76
Peanut, rust on. 132; wilt, 156
Pear. bliRht. 75, 130
Peean, dineane of, 42
Pellicularia, koleroga. 130
Peltier, (J. L., boo Stkvexh, F, L.
Pennington, L. H.. Boleti and myco-
rhtia upon foroBt tnn^ and an unuBual
mycorhiia upon white oak (abBtract),
74
Peony, effect of tar imioko on, 32
Pepper, black spot of. 63; fruit-rot of,
308; Bclerotium diBcaBo of, 64
Peridermium. acicolum. 20. 68; cameum,
66; cerebrum. 315; delicatulum, 68;
harkncBBii. 20. 315; Helianthi, 67; in-
conapicuum, 68; ipomcrfp, 67; Btrobi,
17. 225; terebinthinaceum. 67
Persea, americana. 350
Petunia, 160
Pesiia ciborioidefl. 433
PharbitiB, 68; barbixera. 67; hederacca,
67
PhaaeoluB. vulgaritt. 01
Pheidolo, anaataBBii. 'M\U
Phyllachora, KruniituM. .Vi. 131; grntiB-
Btma, ^i50
PhylloMtictu. hortorum. 7H; ftolitaria. 202
PhyBalift. alkokongi. 161; angulata, 160;
craBBJfnlin. 160; philadolphira. 160
PhyBo|M»llft, vitJH, 132
Phy tophi horn, rnrtorum, 126; crythro-
B<»|»tira. 12«i; inf««j«!an». 122. 262. 319;
terreitria, I'JO
Picea. cngf*liii:innii. 7s. 17«». .'Ci2; oxrolMa.
47; iiiorin<lii. .V)2; ruU'iiM, 47
PiKMKlHKl.. F. J., WM* .STAKyA.V. K. ('.!
FactorM nfforting th<* paniHitiBni of
I'litilaico «oa*. 21^1 .'Ul7
PlEPKk. KhNKHT J.. Hi \IPHIIKY, C .1..
and AiMKF.. S. F.. Synthetic rultun*
nic<iiii ft»r \%*MMl-<li'^trcninpc fungi.
21 f '220
Pierce, Newton B., Biography of, 143
Pierce, Roy G., Albany conference on
white pine bliBter rust, 54; Early diB-
covery of white pine bliBter ruBt in the
United States, 224; see Hpauloino,
Perley
Pine, mistletoe on, 315; rust on, 17, 58,
67, 77, 95; white, 17, 54, 58, 319; yel-
low, 140
Pink root, of onion, 59
PinuB, attenuata, 107, 140; austriaca,
49; banksiana, 107. 137, 140, 315, 451;
caribsa, 67, 68, 315; clausa, 67; con-
torta. 49, 68, 106, 135, 140, 315; coul-
teri, 107, 315; densiflora, 49, 315;
divaricata, 49, 68; echinata, 49, 67,
68; elliotii, 68
Pithecolobium, saman, 132
Plant diseaseB, literature on, 79--81, 152-
154; 22H-234; 3*22-326; 393-398; 45S-
458; Porto Rican, 66, 130
Plasmopara, viticola, 258
Plectodiscella, Piri, 90; veneta, 91
Pluchea. indica, 160
Plum, leaf spot, 75
P(xl blight, on Datura fastuosa, 328; on
Datura stramonium inermis, 328; on
Datura tatula inermis, 328
Polygonum, aviculare, 92; erectUITi, 92;
ramofliBflimum, 92
Polyi>oruB. baBiiapidioilcB, 171 ; berklcyi,
171; frondoBUB. 171; goetzii, 171;
Mylittae. 171; Polyjwrus, 47; rhino-
cerotiH. 171; sacer, 171; Sapurema,
171 ; Bchwrinitzii, 426; sulphureuB, 172;
tulKTaster. 171; umbellatus, 171
PolvBtictUB, abietinus, 380
Poppy, effect of tar smoke on, 32
PopuluB. acuminata, 353; angustifolia,
3.'>.'i; balBamiffTH, 3.5.3; tremuloides,
108. .rvl. .380; trichorarpa. 108. 223, 353,
380
Poria. weirii, 176
PonMliBculuB, iM'n<iuliH. 47
PoHKY. (1. O.. tH'V M<<'riiBiN, W. A.
PotaMitum ryanidr. conduction in
pl.HntM. 443
Potato, curly dwarf, 71 ; effect of tar
Biiioke on. 32; leaf roll, 72; lightning
injury to, 140; malnutrition disease
Index
IX
of, 70; mosaic, 71, 72; necrosis, 71;
powdery scab, 72; Rhizoctonia on, 74,
209; rot, 178 *
Potter, Alden A., and Melchers, Leo
E., Ecological observations on Usti-
lago Zeai (abstract), 73
Pouzolzia, 160
Prunus, avium, 48; ilicifolia, 191
Pseudomonas, Citri, 58; fluorescens, 200;
papulans, 200
Pseudoperonospora, 133
Pseudopeziza, 70
Pseudotsuga, taxifolia, 108, 176, 315, 380,
426
Pteris, 35
Puccinia, Antirrhini, 265; triticina, 224
Puccinia, 18; Canns, 132; glumarum, 73,
142; graminis, 30, 73; graminis tritici,
73; luxuriosa, 93; rubigo-vera, 143;
subnitens, 92
Pucciniastrum, pustulatum, 109
Pucciniopsis, Caricffi, 349
Pycnoporus, cinnabarinus, 47
Pynis, malus, 445
Pythiacystis, citrophthora, 37, 126
Pythium debaryanum, 126, 391
Quamoclit, coccinia, 67
Quercus,'agrifolia, 20; rubra, 107
Radish, lightning injury to, 142
Rankin, W. H., The penetration of for-
eign substances introduced into trees,
^13; The control of white pine blister
rust in small areas (abstract), 58
Rands, R. D., The production of spores
by Altemaria Solani in pure culture,
316; Alternaria on Datura and potato,
327-338
Raspberry, anthracnosc, 83; effect of tar
smoke on, 32
Ravn F. K0LPIN, Jens Ludwig Jensen, 1-4
Razoumofskya, americana, 140; campy-
lopoda, 315; occidentalis abietina, 140
Reedy, C. S., see Jones, L. R.
Reddick, Donald, see Stewart, V. B. ;
see Gladwin, F. E.
Rhamnus, cathartica, 391
Rhizina, inflata, 175
Rhizoctonia, 74; crocorum. 111; micro-
sclerptia, 116; Solani, 111; of the fig,
110; on potato, 209
Rhizopus, nigricans, 178, 391
Rhoads, Arthur S., Some new or little
known hosts for wood-destroying
fungi, 46-48
Ribes, 17, 32, 58, 77; tenuifolium, 372;
nigrum, 449
Ridgwat, Charles S., Methods for the
difTerentiation of pathogenic fungi in
the tissues of the host, 389-391
Rogers, John M. and Earle, F. S., A
simple and effective method of pro-
tecting citrus fruits against stem-end
rot, 361-367
Root rot, of apple, 77, 223
Roripa, palustris, 93
Rosa, 32
Rose, crown canker, 408-417; effect of
tar smoke on, 32
Rose, Dean H., Blister spot of apples
and its relation to a disease of apple
bark, 198-208
Rosenbaum, J., and Shapavalov. M.,
Strains of Rhizoctonia (abstract), 74
Rough-bark or scurfy bark canker, of
apples, 202
Rubus, 32; idaeus var. aculeatissimus, 86;
neglectus, 86; occidentalis, 86
Rudbeckia, 68
Rudolph, Bert A., A new leaf -spot dis-
ease of cherries, 188-197
Rust, 92, 352, 368; on bread fruit, 131;
on canna, 132; on clover, 75; on cot-
ton, 133; on forest trees, 106, 135; on
grapes, 132 ; on grasses, 73 ; on Gondule,
132; on peanut, 132; on Pinus resinosa,
225; on sweet potato, 132; on wheat,
73; white pine blister, 17, 54, 58, 77,
95, 135, 224, 319, 368
Ruth, W. A., see Stevens, F. L. ; see
Brock, VV. S.
Salix, cordata mackensiana, 109
Salpiglossis, sinuata, 161
Salsola, pestifer, 92
Sarcobatus, 92
Scab, on apple, 76, 221 ; on potato, 72
Sclerotium, bataticola, 65, 308
ScHWARZB, C. A., see Blake, M. A., see
Cook, NTel T.
Index
Schixanthus, pinnatufl. 161
Hchixophyllum, rointnunp, 380
HcHNciDEK. Albeht, Further note on a
parasitic saccharomycet-e of the to-
mato. 52-53
ScHKCiNKR, Oswald, see Edson. H. A.
8coleeotrirhuni, graminis, 69
Senecio. 6K
Septoria. Lyro|>orstci, 65; petroselina
var. Apii, 65
Hereh. 423
Sesamuin. orientate, 160
Hhapavalov, M., see Ho8cnbai:m, J.;
Intoxicating broad, Nai'mov, N. A.
(reviews aS4-.WJ
Siicrbakopk, (\ I).. Buckeye rot of
tomato fruit, llD-129
Silphiuin. anKUstuin, 67; asteriscus, 67;
c(>rn|>osituni. 67; dent at um, 67; gla-
hrurit. 67; intcKrifoliuiii. 67; pinnati-
fidurn. 67; trifol latum. 67
Silvenlraad, diHras<' on CofTea, 115
Sisymbrium. altlHuimum. Ik'i
Smith. Clayton <).. Sour rot of lemon
in California. 37-41
Smith. Fum»:\<K P.. we Oiikkly.
KlNKK. fl.
SyiTH. Halph K.. a new apparatus for
aiM*ptir ultrafiltration. 2tK>-2!)3
Smut, oil corn. 73; on oats, .'isi
Snap Iwan. mosaic. 61; Sclerotinia
lib«'rtiaiia oti. M)
Solaiium. rantlincmw*. 6.'>. 163; melon-
Kcna. 157: niKrum. ItU), .'Wl ; tuU'ro-
utiin. 3 J. 157, 17K
S<»lidnKo. r»7
S<M>ty iiifild^. fin (*amcllia. \'X\: on
oraiiic*'. 1<{<'{
S<iphi:t. |:iiiriata. \r2
Sour rot. of ItMiion. 37
S*»y kn'tin b;irt«Tial bliuht of. Ik'*
Spar:iN«in. rn!»pa. 167; hcrlM'^tii. 161»;
laiTiinoM.i. liV'J; ridicats, l(W'i:raiiioHa.l69
Sp\t i,i»iN<.. I'liiu.i.v \ot«H <»n Cronar-
tiuiii Cniiiptotii:!' III. !'.» 51 : K\i<icnn»
of ih«' 'tviT uiiii«rm>: of Cronartium
ribii'ola alKftract . 5s : \«M'dU« runt on
PiTiu- n-finotia. iri5: and PiKiu i.. Hn\
it. S':iti' ami National <|uarantini'«<
:»i:;tjn-f tlir wliit#- |iin«' bli»»t«T ni-f, 3|0
Spilanthes, acmella, 160
Spongospora, subterranea, 72
Sporobolus, airoides, 93
Squash, effect of tar smoke on, 32;
mosaic on, 61
Stakmax, K. C, and Piemeisel, F. J.,
A new strain of Puccinia graminia
(abstract), 73
Stanford, K. E. and Wolf, F. A.,
Studies on Bacterium solanacearum,
155-165
Stanleya, pinnata, 93
Stem rot. of clovers and alfalfa, 432
Stem-end rot. of cirtus fruits. 361
Stercum, frustulosum, 217;spathulatum,
169
Stkvenh, F. L., Noteworthy Porto Hican
Plant diseas(;s (abstract). 66; (ar-
ticle). 1,30-134; and Hi-th. \V. A.,
Pkltikk, (J. L.. and Mall<kh, J. U.,
()l>servations on |M>ar blight in Illinois
(abstract), 75
Stkvknh, Nkil K.. and Hawkins, Lon
A.. S<ime changes prcKluced in straw-
U'rry fruits by Uhizopus nigricans,
17H-184
Stkvknhon, John A.. Lightning injury
to sugar cane. 317; an epiphytotic of
cane <li»eaMe irt Porto Hico. 4JS-425
Stkwaht. F. ('.. \S it(lH's-br<M>m8 on
hickory tn^^'H. IV* IS7
Stkwakt. V. B.. A twig and leaf dis-
fa>M* of Kcrria ja|M»nica. 3t>{^ M)7; The
fM'H'nnation of (Vonartium ribicola on
rurraiit, H1M5(); and UKi>i)irK. Don-
am*. li<*an mosiiic iabHtract>. 61
Stin>ella. Havida. 131
StrawlM»rrv, effect of tar smoke on. 32;
rot of. 17S
Stizolobium, nivcum. H>1
Subulina. o<'tona. 371
Sugar \H*i*\. curly top of. JtiO; light-
ning injury to. Ill
Sugar cam*. ilisiM.***-. lis; leaf spot of.
131; linhtning injury t«». 317
Suntloui-r. elTiTt of tar snn»k«" on. 32
S\Nr«f p<»t:iio. ihan*«»nl rot of, 312;
< '>^to.^|lora batata on. 71; rust on, 132
Svnt-dn'lla, n<MliMt»r:i. Hi()
Index
XI
Tarvia fumes, effect on vegetation, 32
Taubbnhaus, J. J., Two new camphor
diseases in Texas (abstract), 59; and
Johnson, A. D., Pink root, a new root
disease of onions in Texas (abstract),
59
Tatlob, Minnie W., Preliminary re-
port on the vertical distribution of
Fusarium in soil, 374-378
Tea, disenfie of, 132
Tectona, grandis, 160
Thlaspi, arvense, 93
Thomas, H. E., see Fromme, F. D.
Thrinax, ponceana, 131
Thuja, occidentalis, 47
Timothy, Scolecotrichum graminis, 69
TiSDALE, W. H., Relation of temperature
to the growth and infecting power of
Fusarium Lini, 35^360
Tobacco, wilt, 155; Cercospora nico-
tiansB on, 348
Tomato, buckeye rot of, 119; damping-
off of, 319; fruit rot of, 60; leaf-spot of,
327; lightning, injury to, 142 parasitic
saccharomycete of the, 52; Phyto-
phthora infestans on, 319; wilt, 155
Trametes, camea, 380; heteromorpha,
380; hispida, 380; lacerata, 380; peckii,
380; Pini, 176; rubescens, 379; serialis,
380; trogii, 380; variiformis, 380
Trichoderma, lignorum, 424
Trifolium, hybridum, 70, 432; incama-
tum, 70, 432; pratense, 70, 432; repens,
70,432
Triticum, vulgare, 73; compactum, 73
Tropseolum, lobbianum, 161; majus, 160;
peregrinum, 161
Trumbull, H. L., and Hotson, J. W.,
The effect of roentgen and ultraviolet
rays upon fungi, 426-431
Tsuga, canadensis, 46; caroliniana, 109;
heterophylla, 108, 380, 426
Tuberculina, maxima, 139
Turnip, lightning injury to, 142
Tylenchus, tritici, 56, 452
Tyromyces, ca;sius, 47
Uncinula, necator, 66
Uredo, Artocarpi, 131
Uromyces, Arachidis, 132; Dolicholi, 132;
fallens, 70, 75; JaniphsB, 133
Ustilago, Avenge, 381; Isevis, 381; Zeae,
73,294
Venturia, pomi, 221
Verbena, erinoides, 161
Verbesina, 67
Vemonia, 67
Verrucosis, GO
Volutella, fructi, 59
Walker, J. C, Studies upon the an-
thracnose of the onion (abstract), 59
Waite, M. B., Common and scientific
names of plant diseases (abstract), 60
Wandering-jew, effect of tar smoke on,
35
Weir, James R., Notes on wood-de-
stroying fungi which grow on both
coniferous and deciduous trees, II,
379-380; and Hubert, Ernest E.,
Recent cultures of forest tree rusts,
106-109; Pycnial stages of important
forest tree rusts, 135-139; New hosts
for Razoumofskya americana and R.
occidentalis abietina, 140; Sparasis
radicata, an undescribed fungus on
the roots of conifers, 166-177; Note on
Xylari^ polymorpha and X. digitata,
223; Cronartium cerebrum on Pinus
resinosa, 450-451
Western wheat-grass, bacterial disease
of, 225
Wilt, cabbage, 375; flax, 375
Wheat, Tylenchus, tritici, on, 56, 452;
leaf-rust of, 224; rust, 73
Wilt, Bacterial heart, of celery, 64;
crown rot, of celery, 64; of nastur-
tium, 160; of peanut, 156; of tomato
and potato, 155
Witches-broom, on cherry, 185; on hick-
ory, 185
Wolf, F. A., see Stanford, E. E.
Wood-destroying fungi, 46, 214
Xylaria, polymorpha,77,223; digitata,223
Yam, Cercospora carbonacea on, 351
Zantho^ylum, fagara, 339
2iebrina, pendula, 35
Zygosporium, oschioides, 350
KHHATA FOR VOLUMK VII
Pa^o 4i^ line 27, for urtnlinospores read urediniospores.
I'agc 113, 114, and 115, for C(»rticum read Corticium.
Pago 113 lino 9 and page 114 line 4, for ochraUucum read oc/iro/cucum.
Page 132 line 5. for CrphaUurujt read Cephaleuros.
Page 134, for Illinois Tniversity read University of Illinois.
PaKo 1H4 line 7, for frum rea<l from.
Page ia3 line 1, for 30 read 50.
VHge 208, section 6 of Huinniary, line 3, for diseases read diseased.
Page 350. legend for figure 3 should read Zygo»porium anchxoides.
Page 360, 370. and 371, in table headings for uredinospores read urediniospores.
Page 370, table 1, column K, for JG* read 26.
Page .'Wi line 10. for Tomasski read Pomasski.
Page 3110 line 12, strike out comma after water/
Page 418 line 1, after cane insert di.-ease.
' •* , ■^
PHYTOPATHOLOGY
VOLUME VII NUMBER 1
FEBRUARY, 1917
JENS LUDWIG JENSEN
(1836-1904).
F . K0LPIN R A VN
With Portrait, Plate I
The introduction of "hot-water treatment" is one of the most impor-
tant steps forward in practical plant pathology, for it means a new prin-
ciple both from a theoretical as well as from a practical point of view.
Therefore it will surely interest the readers of Phytopathology to be-
come acquainted with the originator of the method, because in it we
find an example of progress due to a man from practical life, who had
scientific qualifications as well, and the ability to carry on independent
research.
Jens Ludwig Jensen was born January 9, 1836, near the little town of
Odder in the Danish province of Jutland, where his father was forester.
In 1855 he passed a normal school examination and later supplemented
his education by studying natural science. He taught school imtil 1872.
In 1868 he started a weekly agricultural magazine, of which he remained
the editor imtil 1880, and from 1879 until his death, he was the publisher
of a widely circulating advertiser.
In 1872 Jensen resigned his position as teacher, and together with a
colleague started a company for selling scientifically tested seed. This
has had a great influence on the development of modern agriculture in
Denmark. He took part in the daily routine of the business until 1881,
and remained one of its directors until 1896.
Jensen never held any official government position but he managed
to arrange his practical affairs in such a way that he could spend much
time in study, and in 1881 he organized this work as a private statistical
institute which he called Bureau Ceres. Its object was a systematic
collecting of observations made in practical agriculture and an experi-
menting both in field and in the laboratory, which he had equipped in
his private home. Although many of Jensen's methods were primitive,
•• .•••• • * » 1 • m #••••• • •
••'•••• •••• % ••*••' '•••*••• •*
• ••• »..•• • ••••,.• ••_••• •••
• ••• ••••••••• ••••*•* •••• •• ••
2 Phytopathology [Vol. 7
yet his work is so exhaustive and original that his investigations of potato-
late-blight fungus (Phytophthora infcsians) and smut in cereals are among
the most important contributions to our knowledge of these diseases.
Jen8<»n*H incentive for investigating the potato disease was the work
done by his company in introducing new varieties of potatoes into Den-
mark. In the years 1878-1881 more than one thousand experiments
were made in all pjirts of the country and wliile inspecting these Jensen
had an opportunity of studying the appearance of the disease under
various conditioius. This suggested infection experiments in which he
studied the spread of the infection from the leaves to the tubers rjid
more particularly the ability of the soil to retain the s|K)resof the fungus
by filtration.
In 1882 Jeii.**en published \\\^ exixjriments which resulted in the develop-
ment of a methoil, "protective moulding/' to prevent the tulxjrs from
infection. He discovered. tcK>, that by postponing digging for two weeks
after the wilting of the top, tul)ers otherwi.se subject to late attacks of
the du^ease might Im» kept from infection.
In June 1882 Jen.*4en U^gan his imi)ortant exi)eriments concerning the
influence of tem|)erature on the development of the fungus. He dis-
covere<l among other things, that this could not grow in a temperature
of under 5°(\ or over 2-r(\ Therefore the storage temjKTature of po-
tatoi^s during the winter should never go liigher than bW These ex-
periments gave Jen.**en the clue to understanding why the fimgus attack
is severe only in temixjnite (*limates, and he formed an hypothesis for
explaining why the iK)tato di.*it*jise was not o!)served in North America
and KurofH* before 1840. First then was the development in the means
of trans|M>rtati<»n such that the |)otatoes from the plateau regions of iSouth
Ameri<*a, the original home of the |K>tato an<l the potato fungus, might
l)e carric<l .•io (|ui('kly through the tropical zone that the hyphae in the
tulK»rs di<l not die on the way.
In ()(*tolMT 1882 Jeimen succee<le<l in demonstrating that the hyphae
an<l s|)on*s which were found in and on the dise:ised tul)ers could \ye killed
by applying a temi)eratun» <»f M)W for four Jiours to the potatoes with-
out affecting their germinating iH)wer. The heat must 1k» applied as hot
air for the germinating |H»wcr sufTered too much when the tubers were
immersed in hot water.
In 18K;J-1H8-1 JeiLK^'u published his method for <lisinfecting seed jx)-
tato«»s by heat. When this was wskhX it was |)o.*isible to prevent the pri-
nuirv attacks of the diM»ase and to delav the se<*ondarv attacks at least
one to two ww»ks.
Until )H>rdeaux mixture ap|M'ared for the first tune in 188G as a preven-
tive of disease. Jeri.Hen's system for fighting the jKitato fungus was the
1917) Ravn: Jens Ludwig Jensen 3
best established method both practically and scientifically. In 1886 his
work received recognition from La Soci6t6 Nationale d' Agriculture de
France, and he was awarded the large gold medal of the society.
In 1885 Jensen began to investigate smut in cereals, and by means of
cross-inoculation experiments he succeeded in demonstrating that the
old well-known species Ustilago segetum should be divided into four varie-
ties which he named, tritici, avence, hordei nuday and hordei tecta. The
distinctiveness of these varieties was later confirmed by the mycological
research of Rostrup, Brefeld, and Kellermann and Swingle. These in-
fection experiments made it appear most probable that smut on barley
and oats was caused by infection during the blossom period and that
the smut spores (or hyphae) are to be found under the glumes in the
ripe kernels. The importance of the r61e played by infected seed was
further demonstrated by experiments proving the impossibility of in-
fection through the soil and manure.
In 1887 and 1888 the most important experiments in disinfecting seed
were published. Jensen compared' the chemical remedies proposed by
others (copper sulphate, sulphuric acid, quicklime, salt) with the hot
water treatment he himself had found. He first applied hot air during
a long period as for the potato fungus, but the results were for the most
part unsatisfactory. However, an experiment made in the summer of
1887 in treating oats with hot water, circa 55°C. for five minutes, resulted
in killing the sniut without affecting the germinating power of the oats.
This experiment forms the starting point for the development of the
"Jensen hot-water treatment."
Jensen now made this interesting observation: Smut in barley is not
to be killed by the same treatment as smut in oats, but if damp barley
seed has been subjected to a temperature of 53®C. for five hours in a corked
bottle the smut disappears. Jensen explained this phenomenon by sup-
posing that a five minutes^ immersion in hot water was insufficient to
moisten the smut germs hidden in the seed and that they therefore were
treated in a dry condition which could not occasion their death. By the
slow heating of wet barley the moisture had time to penetrate and soften
the smut germs and these were therefore killed by the temperature ap-
plied. If this were true Jensen thought that smut on barley could be
prevented by soaking the seed in cold water and then applying the usual
hot-water treatment for five minutes. Experiments made in 1888 proved
the truth of this theory.
As we see, Jensen had now discovered two variations of a method
according to which all forms of smut on cereals might be fought. He
also proved that hot-water treatment could be used for smut on Bromus
and Arrhenatherum and that it might be used as a preventive measure
against certain types of damping-off in sugar beets and mangels.
4 Phytopatholoot [Vol. 7
Jensen's method won and deserved much recognition both in Denmark
and abroad. However, it was a great disappointment to him to see that
many farms which had introduced the method abandoned it later. It was
too complicated for general practical use where no steam was available.
Nor was he successful in his attempt to form a company for building a
factory for applying the hot-water treatment of cereals on a commercial
scale.
After this Jensen discontinued his work with the hot-water treatment
and devoted himself to experiments with chemical remedies. Among
these he preferred potassium sulphide, first tested by Kellermann and
Swingle. This sul)stance was the main ingredient in the so-called Ceres-
powder, manufactured by Jensen and placed on the market in 1895.
The last years of his life until hLs death, August 10, 1904, were largely
devote<l to exjieriments with this remedy and agitation to bring it into
practical use.
During the yean* since Jensen's death the hot-water treatment has
again come to the front. In connection with many of the Danish dairies
and breweries, coojierative institutions have l)een established for dis-
infecting Heed with hot water. During the past year several seed firms
have built factory plants for the hot-water treatment in combination
with a plant for drying the seed. The seed which has been thus treated
is sold with a guarantee for its freedom from smut and leaf-stripe disease.
THE PENETRATION OF FOREIGN SUBSTANCES INTRODUCED
INTO TREES
W. H. Rankin
With One Figure in the Text
Meyer^ in 1808 succeeded in introducing a dyeing liquid into the roots
of a small tree by cutting the stem and immersing the upper part of it
in the liquid. The solution penetrated after some time into all the roots
with the exception of their tips and the slenderest rootlets.
Boucherie* about 1840 patented his method of preserving timber for
building purposes. He made a shallow groove around the tree and
covered it with a belt. The space under the cloth was then connected
with a barrel containing the preserving liquid. The solution was absorbed
and ascended to the branches and leaves. Later, he modified his method.
A canal two centimeters in diameter was made through the stem and
from the latter cuts were made with a saw on both sides as far as possible
without allowing the tree to fall over. The liquid was distributed up and
down the stem. The area saturated, however, decreased rapidly in
breadth in the downward direction. He states that the best seasoit for
thorough penetration by this method is autumn. It is doubtful as to the
meaning of the expression "thorough penetration" since he further states
that if there are hard knots or rotten spots at the base of the tree the whole
strip of wood above them did not become saturated at all and the same
was true with the central part of the core of deciduous trees.
Hartig' introduced colored solutions iiito the growing stems of treed.
He bored two holes at right angles to each other in the trunk and intro-
duced the colored solution into them. It was carried to the top of the
tree but in transverse sections made of the tnink the coloring of the wood
was not uniform. Only those vessels directly above the canals were
colored, forming a cross in the sections.
* Meyer, J. C. F. Naturgetreue Darstellung der Entwickelung, Ausbildung und
des Wachsthums der Pflanzen und der Bewegung und Functionen ihrer Safte. Leip-
zig. 1808.
' Boucherie, M. A. Mdmoire sur la conservation des bois. Annales de chemie
et physyque 74: 113-157. 1840.
Nouvelles recherches siir la conservation des bois. Comptes Rendus 12:
337-339. 1841.
* Hartig, T. (Discussed by Shevyrev 1903: 6-7, but the direct citation to Hartig
is not given.)
6 Phytopathology (Vol. 7
Sachs* in experiments on the rate of ascent of sap in woody plants
used lithium nitrate. He allowed the plants to absorb this through the
roots and found the lithium present at intervals along the stem and at the
tips of the branches. He performed laboratory experiments in which he
showed that such a substance as lithium would progress in the stem almost
as rapidly as water itself, while solutions which dye the cell walls along
its upward coiu-se do not rise nearly so rapidly nor so far, since they are
largely filtered out.
Shev>Tev* was the first to utilize the original negative tension of gases
in the tree as a force for distributing the introduced foreign substance.
He attributes the failiu-e of Hartig and others to get penetration, except
immediately above the incised vessels, to the neglect of this factor. His
method was to attach a funnel or half-funnel to the tree, fill it with water
and then make an opening in the tree with a chisel or auger, underneath
the surfaw of the liquid. As a modification of this he devised a metal
tulx; which wjis jireviously connected with a reservoir containing the feed-
ing solution. This was forced at one end into the bark. The other end
wa.s closed with a rul)l)er stopjx^r through which a bit was inserted. Thus
when the lx>riiig wjis done the solution from the reservoir penetrated
immediately into the wound and the sucking jKJwer due to the negative
tension of the gii.**es in the tree was utilized to pull in the solution. He
states that by this method the dye penneates not only the iierial but
also-the radicate i>jtrts of the plant. .\s to the results, he states: **The
absorl)ed li(|uid luid risen to the top and colored all the veins of the leaves
and even the veiru* of the !>orries on th(» gra|K» vine. The dye could be
det<M'te<l five feet Ik»1ow the surface on all of the roots of the birch, apple
and itsh trees. Thus the first part of the problem is solved; we can in-
tnxluce a licpiid in a <li!!sired quiuitity into all iMirts of a tree. '* Further
he states: '*The vessels iiicise<i in the licjuid, al)«orl>ed and distributed
it to all parts of a living tree. Only the pithy, dead portion of the tree
was not saturated with the li(|uid althougli its absorption by the rays oc-
curred (with an oak). The li(|uid entered the roots as well as the leaves,
twigs and fruit." The data given for ea<*h tree fed do not indicate tliat
he obtaine<l anything more than the satunition of the sap wood and bark
• Sarh?*, J. Kin H«*itrag xur KcnritnifiK livn aurHtcMgendrn Suftntrorii in trun»piri-
nmivu Pflanien. .\rb. lM»t. Innt. \Vuril»urK 2: IIH-IM. 187S.
* Shi'vvrc'v, Ivan. <Kxtrara4lirati» nutrition of <liMeaJ*ni Xrvvn with the aim of
curinic thrni and (i«»j*tn»yinK their paruiiitejf. ) M. Z. and (1. I. Foreatry Dept. Re-
port to Foreatry Department al>out injuriouh iiiHerts, pp. 1-51. 1903. (Roprintinl
from SeUk. Khoi. i. LyetMiv. VMKi: 5H la'M
(Supplement** to the "Fxtraradirate nutrition of <liHoaMe<l trees with the
aim <if runng thefii and <le<«troviriK their (xiriudte?*". i Zemledelt<*heskala Kaiota
(Agrirultufal (iaieti** No- .'{. I. .'>. ft ItKM. ^Reprint eonHuite<l. pp. 1 \li).
1917] Rankin: Penetration op Substances 7
of root and stem and the leaves, for he mentions specifically that the
medullary rajns in the case of a single oak were colored.
Roth,* Goff,' Mangin,^ Mokrzecki,® BoUey,^® Simon," Fron,^ and others
have fed trees with various types of solutions, including colored and
nutritive substances. The majority of them have used nutritive salts or
poisons in anticipation of curing physiologic ailments or inhibiting plant
pathogenes and insects. No accurate data are given on the penetration
except that the solutions in many cases were found to reach the leaves
and some obtained penetration of the roots.
choice of substances for experiment
It seems from the very nature of colored solutions, such as methyl
blue and eosin, that they would not be suitable for determining accurately
the greatest possible penetration obtainable by introducing a foreign
substance. The staining quality is very helpful in tracing the rate and
distance which the substance has advanced, but at the same time much
resistance must be encountered by such substances and finally much of
the original quantity absorbed will be adsorbed, filtered out and chemi-
cally united with the different plant parts which it will stain. The utili-
zation of substances such as the lithium salts overcomes these disadvan-
tages although the actual ascent is not visible. The salts of Uthium are
for the most part soluble in water, they are not used up rapidly in the
metabolic processes of the plant and most important of all the minutest
trace can be detected with the spectroscope. Lithium nitrate in solution
was used m the experiments reported below.
* Roth, Carl. (A method for artificially feeding trees.) Chem. Ztg. 20: 344-
345, fig. 2. 1896.
^ Goff, E. S. The application of artificial root pressure to recently transplanted
trees. Wisconsin Agr. Exp. Sta. Ann. Rept. 14: 272-282, fig. 4. 1897.
*Mangin, L. Sur la nutrition et la defense de la vigne par injection. Jour.
Agr. Prat. 1898: 918-920.
^Mokrzecki, S. A. (A new method of healing and nourishing trees.) Vyest-
vik Tavr. Zenistvo. nos. 11 and 12. 1903.
Uber die innere Therapie der Pflanzen. Zeitschr. Pflanzenkr. 13: 257-265,
fig. 1-5. 1903.
*" Bolley, H. L. (Artificial feeding of trees.) Report of the botanist. North
Dakota Agr. Exp. Sta. Ann. Rept. 14: 42-58. 1903. Ibid 16: 33-65. 1904. Ibid 17:
35. 1906.
" Simon, J. M. (Hypodermic injection in plants.) Jour. Soc. Nat. Hort. France.
(Abs. in Card. Chron. 3:41:8. 1907.)
" Fron, G. (Contributions to the study of the injection of nutrients into fruit
trees.) Jour. Soc. Nat. Hort. France 4: 10: 54^59, fig. 2. 1909.
8 Phytopathology [Vol. 7
forces aiding distribution in the tree
There seem to be three forces which must be depended upon for the
rapid distribution of any foreign substance throughout a tree. (1) By
taking advantage of the negative tension of the gases in a tree in the sum-
mer, when transpiration exceeds the intake of water through the roots,
the solution containing the substance is quickly intromitted. Undoubt-
edly, the currents set up by supplying this ready access to a quantity
of liquid ser\'e to distribute the substance to a certain degree. (2) Most
important of all, however, are the translocating streams of sap in the
tree. The upward movement of raw sap will soon carry the substance
to the leaves and the downward movement of the modified food materials
in the phloem will undoubtedly csLTiy the substance back down to all
parts of the bark. The constant translocation of materials between such
active cells as phloem parench>Tna and medullary ray cells will serve
to distribute it in time throughout these tissues and the downward move-
ment of modified food into the roots would also be expected to ultimately
carry the substance through the root tissues. (3) Except by diffusion,
which is a ver>' slow process, the only movements which can be counted
on to distribute the substance in the heart wood are the translocation
currents in the medullary rays and the alternate withdrawal and renewal
of the water in the heart wood. The water in the center of the tree is
said to act as a reser\'e supply upon which the tree draws during the day
in dry weather. The normal water content of the heart wood is again
restored at night. It wquld apf)ear then that such an oscillation of
currents might 8er\'e evei tually to distribute the substance throughout
the wood.
In other words there is no reason to l)elieve that a foreign sul)stance
intrcnluced into a tree cannot penetrate to all parts provided it possesses
certain properties in itself.
METHOD OP FEEDI.VG
Ten chestnut trees var>'ing from two and one-half to nine inches in
diameter wore fed. The trees were growing in the forest and had small
crowns. Shevyrev*s methods with slight modifications were used in
the«<» ex|)orinH»ntjt. .\ haif-funnel was attached to the tree with putty.
The funnel wiis then fille<l with water and a one-half inch hole was bored
un<l(T the surface of the water with a bmce and hit. The hole was l)ored
to rrach Xhv c<»nter of the tree. .\ one-gallon l)ottle containing the lithium
nitrate* solution had previously lK»en suspended so that the l>otton of the
Ijottlf w'lin slightly hij^her than the hole in the trunk. The solution
was tlu'ii connected with the tree by means of a siphon, made of glass
1917]
Ranxin: Penetration op Substances
9
and rubber tubing, one^iuarter of an inch in diameter. In order to keep
out air and prevent the leaking of the solution the apparatus illustrated
by Rumbold" was used. The rubber siphon was connected to a short
piece of glass tubing inserted through a one-inch rubber stopper. In
attaching the siphon to the tree the solution was first started running and
as the half-funnel was knocked from the tree the rubber stopper with
the glass tubing of the siphon inserted was pressed against the tree so that
it covered the hole. The end of the glass tubing was allowed to pro-
ject into the hole about one inch. The rubber stopper was then held
firmly against the tree by the use of wooden frames and a piece of No. 8
spring-steel wire. Thus the opening was perfectly sealed and after being
once properly adjusted, needed no further attention. The glass siphon
tube, reaching to the bottom of the bottle, was held in place by a loosely
fitting stopper. By slightly raising this stopper a new supply of the solu-
tion could be poured into the bottle.
AMOUNT AND STRENGTH OP SOLUTION PED TO TREES
No attempt was made to keep careful records on the periodicity of the
intake *or to correlate it with any of the factors influencing the rate of
intake. The accompanying table gives such data as were taken. Tree
1 had two days of clear, hot weather on July 15 and 16, when it absorbed
TABLE 1
Amount of lithium nitrate in liters of solution taken up hy chestnut trees
TBBB
I1R8T FBBDINO
BBOOND rSBDINO
THXBD XBDXNO
TOTAL
KUIC-
BBB
0.002 per cent
0.025
percent
0.1 peroent
0.1 percent
Liters
GraniM
Jy. 16-16
Jy. 16
Jy. 16-20
Jy. 90-26
Jy. 26- A. 7
A. 19-0. 10
1
3
2
2
1
2
3
13
3.39
2
2
1.5
Jy. 16-17
1.5
Jy. 17-26
1
2
8
2.45
3
2
i
3.5
2
1
12.5
2.00
4
Jy. 17-18
Jy. 18-26
3
2
5
5.00
5
1
3
2
2
3
11
5.58
6
3
1
4
4.00
7
4
4
4.00
8
2
2
2.00
9
2.5 •
2.5
2.50
" Rumbold, Caroline. Report of the physiologist. Report of the Pennsyl"
vania Chestnut Tree Blight Commission, July 1 to December 31, 1912, pp.45-47»
figs. 39-49. 1913.
Methods of injecting trees. Phytopath. 6: 225-229, pi. 13. 1915.
10 Phytopathology [Vol. 7
five liters of solution in twenty-six hours. Then followed a period of
cloudy and cooler weather which caused a marked decrease in the amount
absorbed by trees 1, 2, 3, and 5 during July 17 to 20. The largest amount
of solution was absorbed inmiediately after attaching. Tree 2 absorbed
two liters the first three hours and tree 3 absorbed five liters the first
nineteen hours. However, the hole in tree 3 reached to decayed heart
wood and the punky wood absorbed an unusual amount. The amount
absorbed diminished rapidly after the first two days and in most cases
practically ceased after the fifth or sixth day. Trees fed the second and
third time did not take in as much as they did the first time. The in-
creased strength of the solution used in the later feedings may hav^ ac-
counted for this. However, no detrimental effect on the tree was ob-
8er\'eil and the one-tenth per cent solution allowed the feeding of a suf-
ficient amount in a shorter time. After the feeding on August 13, all
the trees were allowed to stand until October 10 so that a chance was
affonliHl for more complete distribution. The leaves were just begin-
ning to fall when the trees were cut. A burning of the margins of the
leaves occurred in the case of the smaller trees which had taken up as
much lithium a.s some of the larger trees.
MFrrnoD of analysis of trees
The trees were cut as near to the ground as possible. Cross-sections
about one-half inch thick weie cut from the base, at the point of feed-
ing and ever>' ten feet up the trunk. A few leaves were taken from the
to|>s of the trees. The section.s were then seasoned. To obtain small
block.s from these sections for s[)ectroscopic analysis, a strip al>out one
centimeter wide was sawecl out along the diameter of each section. Where
the bark was thick the cork layer was separated from one end of the strip,
starting with the end which n»pR»sentod a iH)int din^ctly al)ove or lx*low
thr phuT of fet^ding. This >\*as place<l in a viiil and lalM^led. Next the,
inner bark wjis H]>lit off from the wood. Then a small block alx)ut one-
half reiitimeter thick n*pn»senting the sapw(K)d was cut from the strip.
Similar blcM*ks wen* c\it fmm the strip at intervals of al>out two centi-
meters until the sapwcHHl, inner bark and cork was rea(^hed at the other
end (»f the strip. In this way it was considen^d tliat representative por-
tions of the tree wen? obtain(*d for analysis wliich would show rather
accunitely X\u* |)enetration .s<Mnin»d. The l)lo(!ks wen? then incinerated
in ran»fullv clean(»<l rniciblc*s in the bunsen fhiine. The ash was btuned
on a platin\im nc^edle in a colorl(*ss giis fbme and the spectrum observed.
Thr pn»si»nce or alisena* of th<» n»d lithium line indicated whether or not
the lithitim had |x»netnite<l to the \mn of tlie tree represented by the
block U*ing analy.sed.
1917] Rankin: Penetration of Substances 11
RESULTS
The results obtained were practically uniform. The blocks in which
UtUum was found are shown graphicaUy in Ulustmtions 1 to 9 in figure 1.
In all of the trees except 3 and 6 complete penetration of the bark and
sapwood was obtained at and above the point of feeding. In trees 3
and 6, for some reason, the lithium did not penetrate the sapwood on
the side opposite the point of feeding. In all the trees except 1 and 4
the lithium had completely penetrated the sapwood and inner bark of
the sections taken at the surface of the groxmd. In the case of trees
1 and 4 the solution had penetrated the bark and sapwood immediately
below the point of feeding but did not appear in bark and sapwood on
the opposite side. In trees 1 to 6, which varied in diameter from 5 to 9J
inches, the Uthium had penetrated the heart wood only in a few cases,
notably the basal sections of trees 1, 3, 4 and 5 and the section twenty
feet above the point of feeding in the case of tree 2. Tree 4 showed more
heart wood penetrated than any of the others. In this tree the heart
wood was decayed where the basal and breast-high sections were taken
and the bark and sapwood of the basal section on the side opposite to the
point of feeding was not penetrated. However, in the first two sections
alcove the point of feeding the lithium had penetrated for at least two
centimeters inside of the sapwood in sound heart wood. In the case of
the trees of smaller diameter (7, 8 and 9) complete penetration of the en-
tire wood and bark was obtained. These trees (7, 8 and 9) were from
2| to 3 inches in diameter and contained several layers of heart wood.
The leaves and twigs from the very tops of all the trees showed a large
amoimt of lithium present.
conclusions from data obtained
From the above results it may be stated, therefore, that:
1. Lithium nitrate when fed to chestnut trees by Shevyrev's method
penetrates to all places in the tree where there is an active translocation
of food materials, that is, to all parts of the bark and sapwood above
and below the point of feeding.
2. Complete penetration of the heart wood is obtained in trees less
than three inches in diameter. In trees of greater diameter the process
of penetration is slow and does not seemingly follow any definite rule.
Cornell University
Ithaca, New York
12 Phytopatuologt [Vol. 7
EXPLANATION OP PIGURE 1
The fig:ur(« rcfircAont the Atrip8 of wood bewchI out along the diameter of each
croMHtertion rut from tho trees. The blockii analyicd are shown in correct pro-
portion as to siie and position in the strip. Those blocks represented by shaded
areas Rave positive tests for lithium; those represented by white areas contained
no lithium. All the figures are reproduced to a scale which equals one-fourth the
site of the original sections.
The sets of sections are numbered according to the trees which they represent.
The sirctions from each tree are lettered as follows:
.4. Section from base of tree (at ground) •
B. Section at |>oint of feeding (breast high)
C. Section ten feet above B
I). Section ten feet above C
E. Section ten feet al>ovc D
F. Section ten feet al>ovc E
(S. I^*Aves taken from top of tree
When* three blocks are shown at the ends of the strips they represent from the
outside inmard. cork, green bark and sapwood respectively. Where only two are
shown they rrpnttcnt bark and sapwo<Ml.
19171
Rankin: Penetration of Substances
13
inr
I II II II 1
II II II ir~rT
TT-TT
II II II
II ■ II nnn
mnr
n
II II II
0
c
■
A
r
n
II II II
IE
IT
'I 'I II II !•■
e
II II II !■
A
S
n-p-TT
i
II n II
••r
A
S
n
mi
TTTT
n
n
n
IE
TT— n
II II n rr
Fig. 1. Diagrams Illustrating the Penetration of Chestnut Trees
WITH Lithium Nitrate
THE GENERATION OF ALDEHYDES BY FUSARIUM
CUBENSE
Klbert C. Lathrop
The possibility of aldehyde occurrences in cultures of the organism
from the CuImui banana disease Wiis called to the attention of this labora-
tory by Dr. (\ Rumlx)ld localise of the odor accompanying its growth.
The possibility of aldehyde production by the fungus was further suggested
by the work of the author on color production through interaction of
aldehydes and certain plant constituents in connection with the investi-
gation of humus iKxlicH. One of the cliaracteristics of the Cuban banana
disease, whicli according to Smith* is due to the fimgus, Fxisarium atbense,
is the purple, purple-brown, or blackish stain produced in the vascular
bundles of the dise^ised banana plant. This Fusarium also reddens or
purples various culture media. Numerous experiments carried on in
this lalx)rator>' on the action of aldehydes of various chemical constitution
in respect to their effect on plant growth have demonstrate<l that alde-
hydes are uniformly deleterioiLs in action. The generation of aldehydes
by Fusarium cubettM' might therefore account, at least in a measure, for
its pathological action as well.
In reg]ird to the generation of aldehydes by microorganisms Grey' has
shown that acetaldehyde is a product of the action of B. coU cammuntM
on glu<*ose muler anaerobic conditions. That acetaldehyde is a product
of the alcoholic fermentation by yeast was discovered by Roeser,* and
more recently (\ Neul)erg and his co-workers have very fully studied
the mechanism of this reaction. Neuberg and Hildesheimer^ have shown,
for example, tliat acetaldehyde is pro<luced by the action of yeast on
p\Tuvic acid, while NeulM»rg and Kerb' have been able to produce propi-
oni<' aldehyde* by the action of yejist on a-keto but>Tic acid.
Tliat aldehydes are generated during the growth of Fumrium cubenm
on synthetic culture media was experimentally shown in the following
way. Eleven 2-liter Eriemneyer fliisks, each containing nhoui seven
hundred cubic <H»ntimeters of Tschinsky's solution were sterilized in the
» r^ivnce 81: 7.'>l :.V>. VMO.
» HiiK-hem. Jour. 7: :i"»!» 3t«. I*»i:{.
•Ann. Innt. pHMtnir 7: 41. ls<J3.
«Z«Mt. phvHiol. ('h<m. 31: 174. HUl.
•/>>it. phyHiol. ('hem. 47: 413 429. 1012.
1917] Lathrop: Generation op Aldehydes 15
autoclave and inoculated with a pure culture of Fusarium cubenae on
May 28, 1915. The flasks were set a§ide in a dark closet and the Fusarium
was allowed to grow at room temperature until January 21, 1916, at the
end of which time the Fusarium was still growing. The liquid culture
media, which had darkened a httle and which had taken on a slightly
penetrating odor, was filtered from the sediment and growing Fusarium.
The clear filtrate, alkaline in reaction, was sUghtly acidified with dilute
sulfuric acid and the acid Uquid was shaken out a number of times with
ether which had been carefully freed from aldehydes. The aldehydes
were removed from the combined ether extract by shaking with a freshly
prepared, saturated solution of sodium bisulfite. The bisulfite solution
was then acidified with dilute sulfuric acid, the sulfur dioxide removed
by aeration under a long ice cold reflux condensor, and the volatile alde-
hydes, boiling under 75°, were separated from the solution by fractional
distillation, and collected in about fifty cubic centimeters of ice cold
distilled water. The aldehyde fraction so obtained was then treated with
a httle soUd barium carbonate and redistilled in order to hold back any
volatile acids. This distillate was tested for the presence of aldehydes.
A few cubic centimeters of the distillate when treated withSchiflf 's f uch-
sine aldehyde reagent gave a red color inunediately . The distillate reduced
anmioniacal silver nitrate solution, slowly in the cold, and very rapidly
when gently warmed. The odor of the solution was that generally given
by aldehydes, especially the lower aldehydes of the aliphatic series. On
boiling a little of the solution with a strong solution of sodium hydroxide
a pale yellow color was produced which disappeared on longer heating,
ft
and the odor of the solution strongly resembled that of lemon. This
reaction is characteristic of propionic aldehyde, as distinguished from
acetaldehyde or formaldehyde. All attempts to form the phenylhydra-
zone or the p-nitrophenylhydrazone compounds failed, probably owing
to the small quantities of the aldehyde which had been obtained. The
aldehyde in the remaining portion of the' distillate was oxidized by means
of dilute sulfuric acid and potassium permanganate solution to a volatile
fatty acid, which was obtained in amounts too small to be identified by
means of the formation of the metallic salts. By the method of obtain-
ing the aldehyde the fatty acids obtainable by the oxidation with the
acid permanganate mixture are limited to formic, acetic, propionic, butyr-
ic, isobutyric and trimethyl acetic acids. The odors of piu-e hot dilute
solutions of formic, acetic, propionic and butyric acids are so suflSciently
characteristic as to be readily distinguished from each other. Hot dilute
solutions of these acids were then compared with the solution of the un-
known acid and the odor of the pure propionic acid and of the unknown
acid were so exactly similar that they could not be differentiated. This
16 Phytopathology [Vol. 7
would indicate that the volatile acid formed by the oxidation of the al-
dehyde is propionic acid. These reactions show that a volatile aldehyde
is formed during the growth of Fusarium cubense on Uschinsky's solution
and that this aldehyde may be propionic aldehyde, although the amount
of the aldehyde which was obtained was too small to make absolutely
certain its identification as propionic aldehyde. The solution remaining
in the flask after the fractional distillation of the volatile aldehydes gave
no reactions for aldehydes either of the aliphatic or aromatic series.
A badly infected banana stalk received from Trinidad by the Labora-
tory of Plant Pathology was examined for the presence of aldehydes.
The stalk was finely chopped and pressed in a fruit press, and the juice
so obtained was examined by the method given above, but no aldehyde
reactions were obtained.
Since propionic aldehyde is a very volatile compound it is possible
that appreciable quantities were formed during the growth of the Fusa-
rium and escaped from solution. The author had hoped to take up this
question and also establish absolutely the identity of the aldehyde formed,
but circumstances do not permit of this at present. The observations
made seemed of sufficient interest to call to the attention of other workers
on this subject. The author wishes to thank Dr. Caroline Rumlx>ld
and Miss Florence Hedges for their kindness in growing the Fusarium
and for furnishing the diseased and healthy banana stalks.
U. 8. Department op Agriculture
Washington, D. C.
DOES CRONARTIUM RIBICOLA WINTER ON THE
CURRANT?
W. A. McCuBBIN
With One Figure<in the Text
In the literature at the writer's disposal very little mention has been
made of the over-wintering of Cronartium ribicola on the currant.
In a few cases there has been a suspicion of over-wintering on this host
but either the evidence was too meagre to be satisfying, or else other
facts appeared later to explain the circimistances so that the general'
opinion at present favors the entire dependence of the currant stage on
a yearly infection from the pine.
In some of the records, however, there are mentioned puzzling occur-
rences of the rust on ciu'rants, either at long distances from pines, or in
circimistances otherwise so suspicious as to suggest that the fungus might
have passed the winter on the currants themselves. Spaulding (5a) has
mentioned, in connection with the distribution of the Peridermium stage
that judging from analogy with Cronartium CompUmiae, the spores of
which are similar in size and shape to those of P. strobiy it is probable
that the latter would be blown only relatively short distances; but he
records two cases in which no diseased pines were to be found near rusted
currants. In another article the same author (5) lists a reference to an
observation made by Nilsson in 1893, where rusted Ribes were found
over three quarters of a mile from any pines.
Eflforts have been made to settle the question experimentally, both by
planting out badly rusted currants in a disease-free neighborhood after
wintering, and by inoculation with over-wintered spores. So far as is
known the latter method has given only negative results. The former
method was employed by Stewart and Rankin (9) using five hundred
rusted currant plants. None of these developed any rust during the
succeeding summer, and the conclusion is drawn that the fungus rarely,
if ever, over-winters on the currant. In 1914 Spaulding (6), in recording
the negative results from similar experiments with two hundred plants,
says, "The practical conclusion is that Ribes plants do not carry the
fungus over the winter and that an outbreak of this disease on Ribes is
to be attributed to the presence of neighboring white pines which have
the blister rust."
18 Phytopathology [Vol, 7
In a former article (3) the writer has detailed some of the circumstances
of the Ontario outbreak which engendered a suspicion that the rust in
question might have ])as8ed the winter on currants, and in the work with
this disease in 1916 additional evidence has appeared to strengthen this
suspicion. The evidence at present available can not be considered
sufficient to establish the point beyond question, but it is important
enough to be worth careful consideration.
In the discussion following several points which have a bearing on the
question are considered: (1) A hypothesis to account for the various
phenomena observed. (2) Agreement of this hypothesis with known
conditions in other rusts of a similar nature. (3) The general and irregu-
lar appearance of the currant stage over large areas in which there is
reason to believe no pine infections are responsible for the disease. (4)
Special cases, where rust has occurred on currants which are far distant
from any possible source of infection. (5) The occurrence of currant
rust in one instance on two out of four plants in a plantation in which
the same four plants, and these only, were badly diseased in the preced-
ing year. (6) The occurrence of a case of currant rust on plants set out
in a rust-free district in order to test over-wintering.
1. HYPOTHESIS TO ACCOUNT FOR VARIOUS PHENOMENA OBSERVED
On account of the difficulty arising from the loss of all the currant leaves
in the fall, thus separating the fungus from its host, no satisfactory hy-
pothesis has been brought forward to account for suspected cases of hi-
bernation. Spaulding (1911) has suggested the possibility of this hiber-
nation, and has put for\^'ard the idea of a hil>emating mycelium, justify-
ing it by reference to ol)served cases where pustules of Puccinia occurred
on currant shoots. The presence of Cronartium ribicola in such suspicious
locations docs not seem to have been established, although the same
author (5a) records the finding of the telial stage of Cronartium ribicola
on |)etio]es and stipules of Ribes.
Judging from ol)servation8 made during the last two years, the only
hy|)othosis which seems to account for all the ol)serv'ed phenomena in
connection with supi)Osed cases of wintering over, is the hibernation of
the mvcelium in infected buds.
A feature of the disease previously mentioned by the writer (3) has an
im|K)rtant l)earing on this phase of the question: Early and complete
defoliation by the rust, followed by a secondary production of leaves,
due to the premature o|)ening of winter buds. It has often been observed
that such secondarj' leaves are also rusted, even when they are only par-
tially opened, and considering tliat the incul)ation period of the fxmgus
1917] McCubbin: White Pine Blister Rust 19
is from ten days to two weeks or more, it must be evident that infection
can take place very close to the bud stage, perhaps as soon as the bud scales
are parted enough to expose the young leaves within. Now in the case
of a shoot producing secondary leaves in this way, the terminal bud
opens first and makes most growth; the one next below it opens to a
lesser extent; and those farther down exhibit diminishing degrees of ac-
tivity, until towards the base of the shoot buds are foimd which still re-
main in a quite dormant condition. It is true that leaves produced in
this way from buds opening late in the fall are killed by the first severe
frost, and often several of the uppermost buds also perish, but some of
these forced buds, without doubt, are able to survive the winter.
The fact that each shoot producing secondary leaves has its buds ar-
ranged in a series extending from the fully opened condition to the dormant
state, shows that there is ample opportunity for such a favorable com-
bination of circimistances to occur, while the actual presence of the rust
on very young leaves is evidence of the capability of the fungus to infect
at this season of the year. The only step that need be taken outside the
realm of fact concerns the assimiption that an infection can take place
early enough in the development of a bud to still leave it capable of pass-
ing the winter.
The field conditions demanded by the above hypothesis are quite ade-
quate for the purpose. The early defoliation mentioned is general in
some plantations, and in a large percentage of others a smaller or larger
area of plants lose their leaves in midsummer on account of rust starting
from one center. The total number of such cases where secondary foliage
has been produced is many times the number of suspected cases of winter-
ing-over, so that even allowing for a lack of infection in some instances,
dying out of the mycelium in winter, and so forth, there still remain several
times the number of plantations or parts of plantations required to ex-
plain the observed outbreaks of the rust.
In this connection it may be noted that the climatic conditions of the
Niagara Peninsula are extremely mild for the latitude; the autumn is
long and open: in the last two years roses have been in bloom in Novem-
ber; the temperature in winter rarely falls below — 12**F., and there is
an early start of growth in the spring. Under such conditions the per-
sistence of the rust on currant foliage until late in the fall, and its ready
occurrence on the secondary foliage, is not a matter of wonder. In addi-
tion the buds are advanced in this mild fall weather far beyond the stage
at which they usually go into the winter in other localities. Perhaps
the less severe winters might also permit the mycelium to remain alive in
infected tissues in a manner that would be impossible in colder localities.
20 Phytopathology [Vol. 7
2. aqrbbment op hypothesis with analogous cases
The view that the mycelium of the fungus might successfully pass the
winter in currant tissue is open to no a priori objection. What is the
constant habit of the fungus in the pine might well become a temporary
or occasional happening on the other host. In this connection it is scarcely
nece88ar>' to point out the known habits of other rusts under like circum-
stances, but the recent work of Meinecke (4) on Peridermium harknessti
has a peculiar interest here. In summarizing this work he says, ''In Cal-
ifornia Peridermium harknesm and Cr&nartium Qiiercuum are to a high
degree independent of each other;" and again, **Cronartium Quereuum
over-winters on Quercus agrifolia; new urediniospores form in spring
around the old, dead sori on old, living leaves, and infect the young leaves."
He believes that since the crop of new spores is formed around the old
dead spots, that therefore the myceliimi must over-winter in the leaf
tissue. The same over-wintering of the myceliimi has l>een found by
Mains (2) to take place in Coleosporium SolidaginiSy the perfect form of
Peridermium acicolum. In this case the pustules of the fungus were
found to arise in spring in the rosette leaves of the Solidago host, in which
the myceliimi had apparently over-wintered. He was able to prove this
point by sectioning the leaves, and finding therein the rust hyphae in
limited areas.
3 AND 4. APPEARANCE OF CURRANT STAGE WHERE PINE INFECTIONS
DO NOT OCCUR
(hi'ing to the suspicions that arose in 1915 concerning the possibiUty
of over-wintering on the currant, a careful inspection was planned for
lOIt), in onler to <letennine whether ca^es of early infection could be
found, which were either so far away from pines as to preclude the pos-
sibility of pine infection, or which were* close only to pines of small siie,
whosi* fn»<Miom from disease (*ould Ih» al>solutely establLnhed. In general
there arc nuvh nuinlM^rs of large pines scattered over the whole of the
Niagara Pcininsula that, even should cas<»s suspi(*ious of over-wintering
Im* found to occur on curnints, the n(*arness of the other host would ren-
der thcw» c:l*m*s valueless from this |Kiint of view. In two areas, how-
ever, the pin<»s were so few in nmiiUT that an early ap|H»arance of the
curniiit ^tag(* in them would Im* difficult to c^xplain on the gnumd of pine
infection.
One «if thi'H* areiu« (comprises that part of (irantham and Niagara town-
ships included within the d(»tted line on the map (fig. 1), which it will be
nf>ted also n^f^ords the (K)sition f»f all |>ine and currant plantations. In
191') the rust in this area did not start inmi a |)articular (*enter or centers
1917] McCvbbin: White Pine Busteb Rost
I -3
I. i
is -8
22 Phytopathology [Vol. 7
and spread outward therefrom, but it appeared simultaneously and ir-
regularly over the whole district, totally without reference to the few
pines to be found there. In 1916 the very same irregular occurrence was
noted. Ibccept in the Secord case hereinafter mentioned, there were
nowhere any signs which would indicate pine infection; the infection areas
in the plantations of currants were usually few, often only one or two; in
no case was early infection general over a munber of adjacent plantations,
or even plentiful in one; and in most cases the only pines that could be
suspected were more or less surrounded by disease-free currants.
Of the twenty-nine cases of currant rust (plantations) found in the
area indicated, sixteen were so situated as to be regarded with strong
suspicion. Eight of the sixteen are over a^nile distant from any pine
which could possibly be a source of infection, and in all the twenty-nine,
except in the Secord case noted below, the rust started without any ap-
parent reference to the pines in the neighborhood. Eleven of the twenty-
nine cases originated on cither one or two bushes in a plantation, and in
all of these eleven the primary source could still be distinguished on the
early leaves of the shoot. In sixteen other cases of the twenty-nine the
origin frohi a similar small beginning was evident, but owing to lack of
certainty in these cases they are not included as evidence.
Three of the cases occurring in this area deserve special mention on
account of their typical character, and because of the strong evidence
they bring on the question.
\o. /, Lot 17 y Con, 5, Grantham, In this plantation there were present
on July 6, two Imdly diseased bushes, on both of which the early infec-
tion had started on the second leaf of the shoot and had spread from
there over the rest of the bush and to the adjoinitig bushes only. There
are about twenty young pines four hundred feet from these currants^
but they have been examined several times in 1915 and in 1916, and are
all entirely free from the disease. Aside from these the nearest pinee
are over a mile distant with disease-free currants intervening.
So. 2, Lot f). Con, S, Grantham. Here are four old black-currant plants
in a H*)mewluit neglected garden. On July 6 two leaves on one of them
had very old infection spot«, while around these on the bush secondary
unHlincKfnwtules were present. The other bushes were absolutely free
fn)ni the ruj<t. As may l)e MH»n by reference to the map, there are three
lots of pines southeast of thus place, and one northeast. All these are
young pines and are fn»e fn>m any signs of the l)li.ster rust. This disposes
of all the white pines for nearly two miles in every direction except one
forty-yi»anM>ld tree in I>ot KM, Niagara Tp., which is one and a fourth
milc*s distant.
1917] McCubbin: White Pine Blistek Rust 23
No, S, Lot S6y Niagara Tp, The map shows the infected plantation
to be over a mile from any pines whatever. It is moreover in the center
of a currant area of which it alone is rusted. On July 15, the disease was
found at one end of this plantation and the outbreak was traceable to
several very old spots on early leaves.
In the second of the two areas mentioned, which includes the town-
ships of Willoughby, Crowland and Bertie, in Welland Coxmty, the con-
ditions are exactly the same as have just been outUned for the Grantham-
Niagara district. The 1916 infection is irregularly scattered over the
whole territory without any reference to the pines. Out of 185 currant
plantations examined here seventeen were foimd to be diseased, of which
fourteen were suspected to be cases of wintering over. In four of the
cases the currant rust was found from one to two miles from the nearest
pines, and even then these pines were not \mder suspicion, being either
themselves far from a source of infection from currants, or else having
disease-free currants near them.
Although the evidence adduced has been confined to the rust outbreaks
in these limited and favorable districts, it must be understood that the
same conditions prevail in other parts of the peninsula, and it is only the
near presence of so many pines in all these districts which precludes add-
ing a large number of other cases of a like suspicious nature. Unless
either the aeciospores or the uredinospores are carried by the wind to much
greater distances than we are accustomed to think, or than our limited
experiences would indicate, the occiUTence of currant rust in this area
in 1916 is very puzzling on any other hypothesis than that of wintering
over on the currant.
In addition to the above there is another line of evidence which has a
direct bearing on the problem of the origin of the yearly rust outbreak
in the Grantham-Niagara area already mentioned. This evidence is
derived from careful examinations made of cases in the district where
young pines and black currants are growing in close proximity, and of
these the most outstanding instance was on Lot 13, Con. 3, Grantham,
which for convenience of reference is called the Secord case.
On the Pecord farm there was a row of sixteen yoimg white pines planted
along the western boundary. These pines were obtained from a native
wood-lot in the neighborhood, and were planted out in 1910. Rimning
from a lane in the middle of the farm to this row of pines are 175 large
black currant bushes in five rows. On the eastern side of the lane, and
about ninety yards farther north, is a small nursery plot of evergreens,
among which were included about 150 young white pines, planted in
1912, and obtained from the Provincial Forestry plantation in Norfolk
County, where they had been grown from native seed.
24
Phytopathology
[Vol. 7
Although the currants on this farm were badly rusted in 1914 there
was no sign of the disease on either lot of pines that year. In the season
of 1915 the pines were all examined four times (May 5, May 14, June 14,
August 9), but nothing remotely resembling the blister rust was found.
The rust appeared again on the currants in the course of the sununer,
but was late in making its appearance, and possibly came in from else-
where. Another inspection of the young pines was made very early in
the spring of 1916, with the same result as before: no sign of the disease
could lx» seen.
Owing to the close association of the two hosts, and the known occurrence
of the currant stage for at least the two preceding years, special attention
was judged to \ye necessary in this case, and accordingly a further inspec-
tion was made on June 6 and 7. On this occasion there were found a
large number of discolored swellings which were undoubtedly the early
TABLE 1
Total numittr of white pint blister rujtt swellings found June 6, 1916, in the Secord
nurnenj and fence row and the age of growth on which they occurred
TKAR or OBOWTM
NUBSBRT
rSNCB ROW
TOTAL
PBBCKMTAOB
1015
0
V
1
0 5
1914
43
M*
127
71.7
1913
39
10
49
27.7
191J
0
0
0
0
Karli«»r
0
0
0
0
* Somewhat dnuhtful. A pwelling l)el()w the end of a broken-off terminal branch.
* One cif thene prcHlure<i the hliMterK noted above. It waa located at the upper
end of the interntnie developed in 1914.
8tHg(*8 of the blister rust, and which had developed so as to be visible
«inr<» the former visit. Only one case of the blister stage was found:
A .sinall twig near the ground l)ore five small blisters. These had already
shcMJ their sjxires. but were still readily recognizable by the slit-like open-
ings. miinHnts of the |M*ri(iiuni, and by a few remaining spores. The pines
were niitiiitely examined over ever>' part of the stems^ branches and twigs,
and all the swellings collected for study, after which the trees were de-
stn»viM|. The rollertion of swellings was then carefully gone over in the
lalH)ratory, and a record was made of the age of the growth on which they
o<vurre<l. Their position on the tree is given in tabular form below.
Sinc<» the two hosts art» here so closely associated that the currant stage
c<Hild hardly l>e presiMit without causing some infection on the pine, we
are enahlcMi to fonn from this table some conclusion regarding the date
of the first apfx^arance of the disea^* in this particular locality. As no
1917] McCubbin: White Pine Blister Rust 25
pine infections have been found on any growth prior to and including
1912 it seems certain that there was no currant rust here before 1913 at
least. It is, of course, possible that sporidia of 1913 might have infected
twigs of 1911 or 1912, but if so it is hard to believe that all such infections
are still dormant when later pine infections have developed regularly
and vigorously on the same trees. This seems to hint at the absence of
the currant rust here in 1913, but this part of the subject will be considered
later.
Since the currants have been known to be badly rusted in 1914, 1915,
and 1916, it is only reasonable to suppose that plentiful infections have
taken place in each of these years on pines so favorably situated, a point
which is well borne out by the presence of such large numbers of them
on the wood of 1914. These must obviously have arisen after the wood
was formed, and are therefore leferable to the rust of 1914 or 1915. The
above table, however, shows an entire absence of infections on 1915
twigs (save for one very doubtful and abnormal case), and since no swell-
ings or other indications of the disease were seen here in 1915, although
the pines were certainly subject to infection in 1914, it seems fair to con-
clude that during the season after infection the fungus produces in the
twigs no symptoms of a visible nature.
If in addition to this year of dormancy it is assumed that the swell-
ings formed in 1916 will reach the blister stage in 1917, it is probable that
the disease in pine limbs follows a four-year cycle, which may be thus sum-
marized: first year, infection in late siunmeror fall; second year, dormant
period; third year, swelling and discoloration, with possibly pycnospores;
fourth year, production of aecia.
This rule of development is not to be regarded as invariable. In some
cases the dormant period may be extended considerably, and on the
other hand cases are known where blisters have been formed in the second
year after infection instead of the third. Under ordinary circumstances,
however, this cycle is perhaps generally followed.
We may therefore consider the swellings included in the above table
as having originated from infections of either 1914, or both 1913 and 1914.
Since no record exists of the presence or absence of the rust here before
1914, we are unable to state that these pines were not exposed to infec-
tion in 1913. But if they were so exposed and infections resulted, we
are compelled to give an explanation of the sudden development into
the swelling stage of infections of two seasons, simultaneously in the spring
of 1916. If currant rust was present in 1913 it is obvious that every one
of a large number of infections made in that year must have remained
dormant for two years, while on the self-same trees a still larger number
starting in 1914 reached the same stage concurrently, after only one
26 Phytopathology [Vol. 7
year's dormancy. One might imagine that adverse weather conditionfl
might bring about such a pecuhar varation in the dormant period, but it
must be noted that during the spring and summer of 1915 there were
collected at Fonthill, some fifteen miles away, about two hundred pine
infections, in all of which the swelling stage in both old and incipient
cankers was abundantly active. Aside from this one would hardly ex-
pect all these cankers to be affected by weather conditions; some of them
might be retarded but others more favorably situated would have followed
the usual course, and would have appeared as swellings in 1915. It seems
far more reasonable to regard all the swellings in the table as the result
of a single year's infection, that of 1914. According to this view they
have all followed the cycle indicated (with the exception noted), and are
due to produce accia in 1917. It would follow from this, that the cur-
rant rust could hardly have lx»en present on these currants prior to 1914,
the year it was first discovered here.
Since the Secord fann, (fig. 1, -4), is ver>' centrally situated in the
to\^aiship of drantham and in the heart of a region thickly planted with
currants, the al)sence of currant rust here is strongly indicative of its
al)sence in the surrounding neighborhood. Additional evidence on this
point is fiunished by four other cases of a like nature in the same district,
where pines of small size also grew side by side with black currants.
In the first of these, (fig. 1, B)^ a large currant field is 25 yards distant
from a number of young native pines, on which no signs of the disease
could l)e found in 1915, In ()ctolx?r, 1916, there were found on fourteen
of these pines thirty-eight cases of the blister rust swellings. Of these
sixteen were on the growth of 1913 and twenty-two on that of 1914.
None could Ik? found on growth of 1915 and nothing was present on any
wood prior to that of 1913.
In the w»cond case (fig. 1, C) there were two young pines within six
fe<*t of the black currants. In 1916 these also developed swellings for
the first time; of the five seen here two were on wood of 1913 and three
on that of 1914.
In the third case (fig. 1. />), the few young pines were about fifty yards
fn>m a short n)w of black currants. The one case of pine infection which
app<»are<l here for the first time in 1916 yunn on a shoot developed in 1914.
The fourth case (fig. 1, A'), dis<'lose<l only two pine infections on a doien
young trees which were within ten feet of a large but slightly affected
black currant plantation. Ik>th infections wcn^ on pine shoots of 1913
growth. There is some a<lditional evidence here in the fact that this
n)W of pines was transplante<l to its presi^nt situation in the spring of
1914 fn»ni a hollow alM)Ut four hundred yards fnnii these currants. Since
there were no other currants within a quarter of a mile of this hoUow,
1917] McCubbin: White Pine Blistek Rust 27
there is good reason to believe that the pines received their infection after
they were transplanted, that is, in 1914.
In all the four cases just referred to the adjacent currants are known
to have been diseased in 1914. Likewise the pines were in every case
minutely examined in 1915, so that their freedom from disease until the
spring of 1916 is well established. Since in every one of the blister can-
kers found in these foiu' cases the disease has occurred only on wood of
1913 or 1914, and none have been visible till ihe spring of 1916, they agree
in every way with the evidence obtained from the Secord case, and to-
gether with it give a very strong support to the view that the ciurant
rust was not present in the Niagara-Grantham region before 1914.
The bearing of this aonclusion on the question of wintering over is
obvious. If the rust did not appear in this district until 1914, then ac-
cording to the probable life-cycle of the fungus no pine infections could
reach the blister stage and start new currant infection before 1917. It
follows therefore that the currant rust of 1915 and 1916 in the area imder
consideration must be due either to spores carried for long distances by
the wind or to a wintering of the fungus on the currants themselves.
Since this area is about ten miles long by five miles wide, and is distant
from the nearest area of infection (the Fonthill district) from ten to six-
teen miles, any attempt to attribute all these rust outbreaks to wind-
blown spores is full of diflSculty. Aside from this there remains only the
wintering of the fungus on the ciu'rants as an explanation.
Objection may be made to the above reasoning on the ground that in
some instances aecia may be produced the second spring after infection
instead of the third. It is probable that such cases are rare. The single
instance of this sort that was met with in the Secord case was the only
one out of 223 blister cankers collected in these two townships in 1916.
The small branch on which it was found was protected by high grass,
and had the puffed, swollen appearance which normally occurs when a
small limb is stimulated by contact with the moist earth. It was evidently
an abnormal case, and probably had undergone a forced development
by reason of its peculiar conditions. The writer is of the opinion that
such abnormalities would rarely occur in large trees, and as for the small
pines, practically all of these in the two townships have been subjected
to such a scrutiny that the possibility of their playing a part in the yearly
outbreak is utterly out of the question. This is especially apparent when
the extent of the early occurrence of the currant stage is taken into con-
sideration along with the observed rate of spread from the infection started
at the Secord farm by the abnormal blisters just mentioned. In this
case the rust began about June 1, at the ends of the five rows of black
currants. By July 6, it had progressed down the rows for a distance of
28 Phytopathology [Vol. 7
only two hundred feet, and two adjoining plantations distant two hun-
dred and three hundred yards respectively/ were still free from any sign
of infection. By August 21, there was only a slight infection on these
adjacent plantations. It is almost impossible to think that one small
infection of this sort, or even a number of them, could have started such
a widespread infection on currants as our survey disclosed.
Moreover, while an objection of this nature might be vaUd for the
rust epidemic of 1916, it could hardly apply to that of 1915 which was of
the same extent and in the same area. Assuming that the rust entered
this district in 1914 any explanation of the succeeding year's outbreak
on the basis of pine infection would require the production of the blister
stage in the spring of 1915, from a blister canker started by infection dur-
ing the previous autumn. Even admitting the possibility of such pre-
cocious development, the blisters formed in this way must either have
been numerous and widespread, or the spores from one or a few of them
must have been carried from five to ten miles. In the first case our care-
ful examination of young pines must have disclosed some of them, at
least ; and in the second case the general and irregular occurrence of snuill
outbreaks without any recognizable center of infection, together with
the known behavior of the already mentioned outbreak of this nature
on the Secord farm, are quite against any such explanation.
5. RECURRENCE ON SAME INDIVIDUAL
The hypothesis advanced al)ove is capable of being tested to some ex-
tent by field ol)8cr\'ations. If the rust winters in buds forced into late
growth by premature defoliation, then it should be possible toestablish
a connection l)etween suspicious early outbreaks of the rust and the
plants or small areas which were defoliated during the preceding summer;
if these areas showing force<l growth were marked in the fall and rust
appeared in them in spring in a largcT percentage of cases than in the
or<linar>' parts of the field, the case for over-wintering would be very
stmng indwMl. Such field work would necessarily have to be done in a
district when» the n^sults would not be interfered with by the presence
of too many pines.
rnfortunately no «yHtematic effort has yet l)een made along this line.
The Humll amount of €»vi(leiire now in hand conies from three plantations
which were niarkecl as defoliated in 1915. Out of the three, two were
subject to early outbreaks of the rust in 1910. while the thin! was free.
In U)th of the two cjises the infection wjis small and startec! from one or
two renters within the defoliatcNl portion. \ |)erhaps more significant
case came up in the field work of one of the ins|K>ctors. In one large
1917] McCubbin: White Pine Blister Rust 29
black currant plantation the inspector who covered the ground in 1915
reported only four bushes, adjacent in a row, as badly rusted, with but
slight infection elsewhere on adjoining plants. These four bushes were
seen at that time by the owner, and when the inspector of 1916 called here,
the owner told him where to find these bushes, which were on the side
of the plantation opposite a small tree. The only case of rust which
could be found in this plantation was on one of the four plants mentioned.
It is conceivable that spores blown from some other place might start
an infection in this one spot, and there only, in two successive years,
but it is so utterly improbable that one can scarcely avoid turning to the
wintering-over hypothesis for an adequate explanation of the case.
6. RUST ON transplants IN A RU8T-PREB DISTRICT
In a former article (3) mention has been made of a rust outbreak which
occurred in a small plot of black currants set out in the spring of 1915
to test for hibernation. The one hundred currant bushes used were all
badly rusted in 1914. They were divided into five lots, of which two
were well sprayed with lime-sulphur, two were left unsprayed, and one
was exposed to infection from rusted currant leaves wintered out-of-doors
and suspended among the foliage in loose wire baskets. A locality was
chosen for the experiment far away from any known rust area; this dis-
trict had few pines and the freedom from rust of the few existing currants
was ascertained during the fall of 1914.
The one case of rust which developed on these plants was on one of the
sprayed plots. At the time of examination, October 18, it was still of
very small extent, involving only one shoot of a single plant. There was
a small original rust spot surrounded by about twenty others of more
recent date.
Owing to the limited extent of the rust here so late in the season there
was some hesitation in attributing it to a wintering of the fungus, and
every other possible source of infection was given due consideration.
The only one of these possible sources that had any degree of probability
was the carrying of aeciospores to this place in an inspection visit made on
May 24. At this date no currant rust had been met with but some inocula-
tions with the aeciospores 6ad been made in the laboratory on May 20.
The writer did not do this work himself but was in the room at the time,
and a few air-borne spores might have adhered to his clothing and have
been thus carried to the field in question four days later. Improbable
as this suggestion may seem, it is the only explanation on the basis Qf
accidental infection which seems to be worth consideration,
30 Phytopathology [Vol. 7
On the other hand the wintering of the fungus on the curranta them-
selves readily explains the case. The fact that the plants were well
sprayed in spring is in perfect harmony with the hypothesis advanced.
Even the late appearance and small spread of the infection, which would
appear to be incompatible with wintering-over, need present no difficulty;
it is well known that while Puccinia graminis may live independently
of the barberry for indefinite periods, yet by undergoing its proper stage
on this host its virulency on cereal hosts is much increased. It should
not l)e too much to expect, then, that in the case of Cronariium ribicola
the second successive seasonal generation on the currant should lack
somewhat of the vigor it would possess after coming fresh from the pine.
The results of the sur\'ey work in the Niagara Peninsula seem to hint
at a confirmation of this view. A great many of the cases of currant
rust observed here during the last two years have been in the type in-
dicated : an old but very feeble rust center, which by the end of the sum-
mer had involved only a single bush, or at most a few adjacent bushes.
Other explanations of this seeming lack of vigor are no doubt possible,
but such a decadence is quite compatible with a hibernation hypothesis.
In concluding this discussion it is clearly recognized that the evidence
submitted is inadequate to establish the point under consideration, but
on the other hand it is considere<l that enough evidence has been adduced
to warrant a strong suspicion of currant hibernation, and this suspicion
holds even though the hypothesis tentatively put forward should prove
to l>e untenable in the light of later investigation. In any case the evi-
dence obtauiod 8er\'es to narrow the field of inquiry to a great extent;
for the conditions outlined above are such that the question of hil>ernation
clearly hinges on the distance aeciospores or uredinospores can l)e car-
ried by the wind; if only for a mile or two, then wintering on the currant
has almost certainly taken place in the area under consideration; if on
the other hand the spores are Iwrne eight or ten miles or farther, another
explanation of the situation in this district l>ecomes easily possible, al-
though the question of hibeniation is not even then altogether disposed
of. In the al)sence of definite infonnation concerning si>ore dispersal
the cjuestion must remain o|H»n until a l>ody of trustworthy evidence can
Ih» acnmuilated on this |)oint, or until more direct evidence is available
on other phases of the subject.
Dominion Fikld Lahoratory of Plant Pathoukjy
St. Catharines, Ontario
1917] McCubbin: White Pine Blister Rust 31
BIBLIOGRAPHY
(1) HowiTT, J. E., and McCubbin, W. A. An outbreak of white pine blister rust
in Ontario. Phytopath. 6: 182-185. 1916.
(2) Mains, E. B. The wintering of Coleosporium Solidaginis. Phytopath. 6:
371. 1916.
(3) McCubbin, W. A. Notes on white pine blister rust. Rept. Dominion Bota-
nist (Ottawa, Canada) 1915-16:
(4) Meinecke, E. p. Peridermium harknessii and Cronartium Quercuum. Phy-
topath. 6: 225-240. 1916.
(6) Spauldino, Perlet. The blister rust of white j}ine. U. S. D. A., Bur. PI.
Ind. Bull. 206: 1911.
(5a) The present status of the white pine blister rust. U. S. Bur. Plant Ind.
Circ. 129: 15. 1913.
(6) Notes on the white pine blister rust. Phytopath. 4: 41. 1914.
(7) New facts concerning the white pine blister rust. U. S. D. A., Bui. 116:
4-5. 1914.
(8) Stewart, F. C. An outbreak of European currant rust. New York (Geneva)
Agr. Exp. Sta., Tech. Bui. 2. 1906.
(9) Stewart, F. C, and Rankin, W. H. Can Cronartium ribicola over-winter on
the currant? Phytopath., 4: 43. 1914.
THE INJURIOUS EFFECTS OF TARVIA FUMES ON
VEGETATION
A. H. Chivers
The following article contains a brief description of the destructive
efToct of tar smoke on plants which the writer had occasion to study
during the sununer of 1914, together with a brief account of experiments
carried on since that time in the laboratory, and under controlled con-
ditions.
The tar compound which was in use for building purposes, and which
seriously affected the neighboring vegetation, is sold under the trade
name, tarvia. For melting the compound a single kettle was set up
alK)Ut sixty feet distant from the nearest corner, and three hundred feet
distant from the farthest corner, of a garden which covered an area roughly
a hundre<l and eighty feet square, and which contained both ornamental
and cn)p plants.
A strong and constant wind carried the fumes over the garden for about
four hours on the afternoon of August 10 and throughout the forenoon
of August 11, during which tune the leaves and stems of the planta l)e-
came coated with a sulwtance of a greasy nature.
The sulisetjuent destruction of the plants was ver>' rapid. The leaves
scHin curled and shrivelled, dried out and fell. At least twenty species
and numy varieties were affect €»(!. Poppies, P, soinnifirum Linn., P, rhoeas
Linn., P, orieuUile Linn., candytuft, Ilxrin amara Linn.; marigolds, Caleri-
dula officinaliH Linn.; azaleas, Azalea vinami Linn.; sunflowers, Helian-
thuM anfiuuM Linn.; strawln^rries, Fragaria in varieties, and squashes,
i^iCurhiUi jH}H) Linn., and (\ maxima Duchesn(», were killed. Pae<mies,
Parouia in varieties, were killed to the surface ui the soil. Roses, Rom
in varieties: hoiiey.^uckles, Ltnicrra Uirtarica in varietit*s; currants,
Wi7x\K in vari«'ti<\s; ra.*<plM'rrieH. black ra.*<pl Harries and blacklnmes, Kubus
in varii'tirs, were dcfoliatet), uimI in son:e instances were killed. Pota-
toes. Snlauum tulMroftum Linn., which occujMed the greater part of the
garden, were stunt€»<l and the yield was greatly reduced. All perennials
showed the effects of the injury in the following season's growth.
IIHIKF Sl'MMARY OF MTKIiATlKE
( )bs«Tvations on the effect of vaiK)r an<l dust fnmi tarred roads, and
smoke from melting tar on neighlK)ring vegetation, have led to extensive
1917] Chivers: Eppbcts op Tarvia Fumes 33
investigation of many tar compounds, and it has been found that while
these compounds vary widely, the smoke and fumes from these are in
general injurious to plants.
Gatin^ investigated different substances used for the surface treat-
ment of roads and his results tended to show that many trees, shrubs,
garden plants and flowers suffered injury from the fumes given off by the
tar, and also from the dust arisinjg from the treated roads. The injury
seemed to be proportional to the distance from the road, the amount
of phenol in the compoimd, and the isolation of the plants. The effect
was shown in the fading of the leaves which were spotted and blackened.
The cells were plasmolyzed and the chlorophyll disappeared. Marked
differences in resistance to injmry on the part of some plants were noted.
Gatin and Fluteaux^ found as a result of studies on leaves and branches
of catalpa and locust that plants which have been submitted diu'ing the
season to the dust from tarred roads had become considerably modified
in respect to anatomical structure.
Mirande^ made a study of the influence of the tarring of roads on plants,
and concluded that the injmy was done by vapors given off in consider-
able abundance diu-ing dry, hot weather. He stated that if trees and
ornamental plants in cities are to be preserved the use of tar on roads
should be made with care. The same author* investigated the effects
on plants of a number of commercial products such as Carbonyle, Car-
bolineum and Carboneine, all of which contained creosote, and a nimiber
of which were used as insecticides. They were more or less injurious,
causing the destruction of the green cells. He urged care in their
application.
Griffon,* as a result of laboratory work covering three seasons, confirmed
the conclusions of Mirande regarding the injurious nature of gases given
off from tar when used in coating roads. From extensive observations
^ Gatin^ C. L. The efifects of tarring roads on the growth of trees in the Bois de
Boulogn. Compt. Rend. Acad. Sci. Paris. 168: 202-204. 1911.
The experimental reproduction of the injury to plants by the vapors and
dust arising from tarred roads. Compt. Rend. Acad. Sci. Paris. 168: 688^90.
1911.
The tarring of roads and its effect on the neighboring vegetation. Ann.
Sci. Nat. Hot., ser. 9, 16: 165-252. 1912.
' Gatin, C. L. and Fluteaux. Anatomical modifications produced on plants
by dust from tarred roads. Compt. Rend. Acad. Sci. Paris. 168: 1020-1021. 1911.
* Mirande, M. The effect of tarring roads on plants. Compt. Rend. Acad.
Sci. Paris. 161:949-952. 1910.
* Mirande, M. The effect on plants of certain substances extracted from coal
tar. Compt. Rend. Acad. Sci. Paris. 162: 204-206. 1911.
* Griffon, £. The influence of tarring roads on neighboring vegetation. Compt.
Rend. Acad. Sci. Paris. 161: 1070-1073. 1910.
34 Phytopathology [Voi^ 7
he concluded that the probable injury to vegetation in the open coun-
try would be small.
CMaussen* exposed plants to vapors of several coniniercial tars, and
found that the various kinds of tar sold for building highways differed
widely as to their effects on plants. The nature and extent of the in-
jury' were closely related to the concentration, thus depending on vola-
tility and temperature, and that species of plants differed widely as to
their susceptibility to the vapors. Certain recommendations were made
in respect to the proper handling of such products.
Gabnay' gave a brief account of the injury to trees by tar used on the
trunks as protection against the ascent of caterpillars, which involved
not only the cambium but also the sap wood, and extended beyond the
limits of the tarred areas. The injury was ascril)ed to the exclusion of
air and the action of acids and salts.
Ewert' investigated the injiu^ to vegetation by smoke-borne products,
and reported a peculiar lacquered appearance on the upper surface of
leaves of a number of economic plants, frecjuent rolling and crumbling
of the laminae, and discoloration over part or all of the surface. Fruits
and garden produce in such neighborhoods showed the effects. Controlled
experiments showed that injury depended not alone upon the amount
of material present in the atmosphere, but also ujwn the heat, dryness
and isolation.
A cas<* most similar to the one under discussion is recorded by Moore*
as having occurred at Woods Hole, Massachusetts, when a collection of
valuable roses was seriously damaged by smoke which resulted from the
burning of a tar and gravel roof in the vicinity of the garden. The effect
of the smoke began to be noticeable during the third day of the fire, and
was indicated not only externally by the falling of the leaves and the
scarring and marking of the young and tender stems, but also internally
where large areas of growing tissue died and the contents of the cells
were shrunken and di**torte<l, the green coloring matter luiving been
completely <lis<irgunize<l. All plants were affected, some were killiHl
outright, and others so weakened that they l)i»ciune much more susceptible
to the attack of fungous db<ea«es.
• ClaiiHHon, r. TUv infliienr<> of tar, particularly that of tarred strccta upon
vegetation. Arl». KaiM. Hiol An«t. I^nd. u. For«tw. 8: 403-.')l4. 1913.
' Cialinay, F. von. The patholoxical action of tar on plants. (Vntbl. Omam.
ForMtw. S9: 497 .VM. 1913.
• Kwert, H. Injury to vegetation by coal tar and other vapors, and protection
therefrom. Zeitiirhr Pflani«»nk U: 257 273, 321 34(). 1914.
• Moore. (;. T. Ho»t!J« vii. Kailroadw. Kho<!ora 5: 93-96 1903.
iai7] CnrvERs: Effects of Tarvia Fumes 35
The very complete bibliography of McClelland^ also should be con-
sulted in this connection.
RESULTS of experiments CONDUCTED IN THE LABORATORY
It was soon found that the injury to plants by tarvia could be dupli-
cated easily in the laboratory, and experiments have been made for the
purpose of determining whether or not what seemed to be facts at the
time of the accident would appear imder controlled conditions. For this
work begonias, Begonia in varieties; ferns, Adiantiun, Aspidiiun and
Pteris; wandering jew, Zebrina pendida Schnizl. and Commelina nvdi-
flora Linn.; and geraniums. Geranium in varieties, were used.
In a comparatively short time after the plants were placed in the path
of the fumes, the same greasy covering of condensed volatile substances
which collected on the garden plants began to appear over the plant
surfaces. Plants three to four feet distant from the source of the fumes
showed an appreciable covering in about three hours.
The symptoms of injury were found to vary appreciably with the species.
Leaves of begonias showed a characteristic sinking of the upper epidermis,
at first in small, isolated areas, which gave a peculiar pocked appear-
ance to the leaves. The pock marks gradually became confluent, and the
entire area lost chlorophyll and turned brown. In the youngest leaves
the first symptoms appeared as yellow spots, three to six millimeters in
diameter, which when examined, were foimd in each case to be an injured
area immediately surrounding a multicellular gland. Older leaves turned
yellow over their entire surfaces and fell from the stem.
Ferns treated with the fumes withered and dried as if subjected to
extreme heat. Geraniums showed a tendency of spotting. In general,
however, the lower and older leaves turned yellow, those of medium age
turned dark brown over the entire surface, while the youngest and only
partially unfolded ones showed dark brown zones on their margins.
Experiments were performed with the purpose of determining whether
or not the injury was due to the interchange of gases through the stomata.
Species of begonias were particularly desirable for these experiments,
since stomata are found only on the imder surfaces of the leaves. Plants
with a single stem bearing about ten leaves were used. The stem was
wound with cotton and then with waxed paper. Some leaves were left
unprotected. For other leaves cork masks were cut to fit the upper and
under sides, the center of the upper mask having been cut away until only
a narrow rim remained. These masks were then pinned in place so that
the leaves were entirely protected on the stomatal surfaces, but exposed
^^ McClelland, E. H. Bibliography of smoke and smoke prevention. Mellon
Inst. Indus. Research. Bui. 2: 1-164. 1913.
36 Phttopathologt [Vol. 7
on the upper Rurfaces with the exception of a narrow margin. The
plants were placed so that the exposed surfaces faced the fumes. In
all cases the injury was as marked and of the same nature as in the un-
masked leaves.
An experiment was tried of painting onto the siirfaces of the leaves with
a camel's hair brush the volatile matter which condensed on the surface
of the glass above the emanating fumes. It made little difference whether
it was applied to the upper or lower surface of the leaf. The affected
areas showed the same symptoms as those treated with the fumes. The
painted spots l)ecame brown and finally dried and dead.
It was assumed from the first that the injury in the garden was due en-
tirely to the effect of fumes on al>ove-ground parts. To confirm this
assiunption, however, the potii were either wrapped in several layers of
paraffined paper, or coated with paraffin and their tops covered with
waxed paper. With plants thus protected the results recorded above
were obtained.
SUMMARY
The results may \yo sunmiarized as follows:
1. The fumes from the compound known as tarvia are highly injurious
to vegetation.
2. liCaves whose surfa<*es were painted with the oily matter which col-
lected on a ccH)l gliiw plate over the emanating fumes showed the same
symptoms of injury as did those treated with the fumes. This, together
with other evidenr<% indi(*ated that the injury was due in large part at
IcNist to the c<»nstituents of the volatile sul)stances which condensed in
the fonn of an oily c*oating on the surfaces of the plants.
3. Plants with no stomata on the up|M»r surfaces of their leaves were
pnitected in resjKTt to all other surfac€»s, and so placed that only the
up|M»r leaf surface was subjected to the fumes. Such plants showed
injury r»f exactly the s<mie nature as did tluise with unprotected stoniatal
surface's. This indinited that the injury did not involve, to any extent
at least, the piissiige of gases through stomata.
4. In a ."iufficient numlxT of ex]M'rimcnts the soil and under-gnmnd
structures were protected from the fumes, showing that the injury* wa.«*
due t«> the action of the fumes on aerial fmrts.
'). The injury varied with the distaixr from the escaping fumes, the
teiii|N*rature of the melting tar, the age of the plant structures, and the
^|M'cies used.
DaKTMOCTII (^oLLI-^iK,
ll-Wdvwi, N. H.
SOUR ROT OF LEMON IN CALIFORNIA*
Clayton O. Smith
With Two Figures in the Text
The fungus causing the decay described in this paper was first iso-
lated some years ago from lemons which were originally infected with
the brown rot fungus Pythiacystis ciirophthora. Artificial inoculations
made at that time on green lemons with a pure culture of the organism
gave negative results, and no further attention was given to it until in
1915. The method of development, morphology and general charac-
teristics of the fungus, noted at that time, were similar to those described
in the present study. The decay has been found in many of the lemon
packing houses of ("alifornia and probably occurs more or less in all of
them. The fungus is not known to have caused serious losses until the
summer of 1915, when the unusually large crop of lemons made it neces-
sary to hold large amounts of fruit in storage for a longer period of time
than is customary, during which time considerable loss occurred in the
packing houses and in transportation. It seemed to be especially in-
fectious with fruit picked in the spring, being most commonly reported
in May fruit. The fungus has also been found causing a rot of Valencia
oranges in transit.
Several popular terms have been applied to this decay such as sour
rot, slimy rot, watery rot. These terms are descriptive of different stages
of the decay. The peculiar sour odor is so constant a characteristic and
one so distinct from those of other decays of citrus fruits that the name
sour rot is suggested for this decay.
Sour rot is a soft decay, during storage, of citrus fruit, especially of
lemons. The tissue when infected quickly softens, but for some time may
retain nearly its normal shape. It, however, changes to a straw color*,
later collapses, becoming more or less slimy with age, and at last is almost
completely changed into a watery mass, which in the packing house often
drips down into the lower fruit of the stacks. Because of these charac-
teristics the grading and sorting of the fruit is ver>^ disagreeable. The
softened areas of the fruit do not at first show any noticeable aerial my-
* Paper No. 38, Citrus Experiment Station, University of California, Riverside,
Calif.
^ Dauthenay, Henri. Repetoire dc Couleurs, p. 31, No. 3.
1917] Smith: Sour Rot of Lemon 39
nally described by Ferraris^ as Oidium citri-aurantn, Saccardo and Sydow
later transferred the fungus to the^enus Oospora. Cultures of the soiu" rot
fungus were submitted to Professor David R. Sumstine, Peabody High
School, Pittsburgh, Pennsylvania, for identification, and he regards this
species as belonging to the genus Oosporoidea,' a group of fungi that is
now separated by some systematists from Oospora because of the slight
differentiation between the mycelium and the sporophores. The fungus
should now probably be called Oosporoidea citri-aurantii (Ferraris), but
for the present will be designated as Oospora. The aerial mycelium of
the sour rot readily separates into spores when moimted in water for
examination, as does also the mycehum growing on the substratum.
Sporophores and chains of spores are with difficulty distinguished from
the myceliimi.
Ferraris* foimd from his study and inoculations, that the fimgus Oospora
cUri-aurantti caused an infectious soft decay of oranges. The individual
points of infection increased in size and coalesced. A strong odor of
fermented juice and a disagreeable taste of the fruit accompanied the
decay. No aerial mycelial growth was at first visible, but under favor-
able conditions, a short very white, wrinkled myceUum developed, form-
ing in contact with the substratum a gelatinous layer. The myceUum
has a constant diameter of about 7 m being described as being perfectly
yellow and granular when growing in the orange tissue. The color of
the mycelium of the sour-rot fungus as observed under the microscope
when taken from artificially inoculated oranges, shows a slightly yellow-
ish color but could hardly be said to be perfectly yeUow. The size of the
conidia as given agrees very closely with that of the sour-rot fungus.
Ferraris recorded the size of cyUndrical conidia as 13.5-19 x 7-7.5 m;
oval conidia 9-12 x 7.5 m; spherical conidia about 12 /z.
Ferraris refers to certain other closely related fungi, causing rots of
citrus fruits. Among these are Oidium fasciculata Berk., probably synony-
mous with Oospora fasciculata Sacc. et Vogl. and Acrosporium fascicular
turn Grev., which is said to occur in decaying citrus fruits in Belgium,
Great Britain, Italy and North America. This fungus differs from Oospora
citri-auraniii in that the mycelium is at first white but changes to glaucus
with age. Oidium tigitaninum was described* from CaUfornia as a pow-
* Ferraris, T. Di un nuovi ifomicete parassita nei frutti di arancio. Malpighia
IS:. 1900.
* Sumstine, D. R. Studies in North American Hj'phomycetes. The tribe Oospo-
reae. Mycol. 6 : 45-61 . 1913.
* Ferraris, T. Loc. cit.
•Carter, C. M. A powdery mildew of Citrus. Phytopath. 5: 193-196. 1915.
40
Phytopatholoqt
[Vol. 7
dery mildew of Dancy tangerine. It diffcnt in size, in shape of span*
Bttd in other morphological characters from tho fundus under oonsideration.
Ootpara cilri'OUTantii is closely rolateil to Oon/jom fcidia morphologi-
cally, a fact fully rectqpiized by Ferrarip, who found similarity in conidia
c4
c?'*
.\. It. r. :ii-n:il mv. .-Iiuni ->l...uiiiii chain- -f •,
into n-H- (h:it fill.. <■'.), a- !.|...r.-^ K. .-..ni.liii. -Ii.
lion F. mI.I I'li^i ■'lioviMK ..11 k1..I.<iI.'v
Ki-niiiiiiiliiiii. li\p!i:il l.niTn-liitin. iitui thr mr
i-iuiiii^ ;iri- f<>titi.-<l. Il<- :il-^i mi'iili<>ti'< lli<- i
wliicii tliov i»i. tiiUKi site ztl.lf to priMhict-. i-
r in U'hirli ih<> omidiii!
niiii'^ nuiiitxT of (tmiilia
i:iltv whi'Ti ihf niv«*elituii
1917] Smith: Sour Rot of Lemon 41
itself breaks up into spores. Certain differences have, however, been
found between these two fungi.
Oospora lactis is probably a composite species into which a number of
closely allied fungi have been placed by systematists. A culture of Oospora
ladiSy with which the sour-rot fungus was compared, was furnished by Mrs.
Flora W. Patterson. This culture was isolated from oysters. A culture
isolated from the same source was also received from Dr. Charles Thom.
Artificial inoculations w4th Oospora lactis from these two cultures and with
the sour-rot fungus on citrus fruits showed that Oospora citri-aurantii ia
pathogenic, causing the fruit to begin to decay within a few days. Oospora
lactis at first produces some mycelial growth on the injured tissues, but
the myceliimi does not appear to be able to attack the tissue adjacent to
the injury and no actual decay takes place. These experiments were
performed several diflFerent times in moist chambers on Eureka lemon.
Navel orange and Dancy tangerine. No infection took place, although
duplicate experiments on the same kinds of fruit with the sour-rot fungus
gave positive results.
The reaction of Utmus milk with the two fungi differs. The sour-rot
organism caused no change in the reaction and probably made but slight
growth. Oospora lactis showed an acid reaction and clearing of mediimi
without separation of the casein.
The spores of the sour-rot fungus appear more regular in size, and more
cylindrical than those of Oospora lactis.
Artificial inoculations were made by puncture with cultures of Oospora
citri-aurantii on the following citrus fruits in moist chamber: lemons,
oranges, grapefruit and tangerine. Positive results were secured in
moist chamber, the rot beginning to show in forty-eight hours. The ripe
or nearly mature fruit is more readily infected. Failure to infect the
green fruit of lemons has frequently occurred. Inoculations on the twigs
of a Eureka lemon gave negative results. Lemons showing the initial
stages of the brown-rot fungus, Pythiacystis citrophthora^ were atomized
with a suspension of spores of the sour-rot fungus. Infection took place
quickly in the brown rot areas and continued to increase as the former
decay advanced. Eventually the surface of the lemons was coated with
the sour-rot fungus. Sound fruit when inoculated with an atomized
suspension of spores or when soaked for twenty-four hours in spore-laden
water were but rarely infected and then probably only in some superficial
injury. Infection of lemons with the sour-rot fungus evidently only takes
place through some injury or from contact with infected fruit.
WnimER, California
A DISEASE OF PECAN CATKINS
H. H. H I G G I N 8
With Two FiorREs in the Text
During the latter part of April, 1916, the writer's attention was called
to an ahnonnality of the catkins (staininate) of pecans, Carya iUinoensiSf
on the ExpcTinient Station plats. Some, or in many cases, all of the
flowers of a (*atkin were slightly distorted and of a paler green hue. The
stamens and inner surfa(*e of the subtending bract were covered with a
white sutjstance whi<'h at first glance gave the impression of white fly,
but which on examination was found to l>e the white spore-cluster and basi-
dia of a fungus l)eIonging to the genus Microstroma. A little later when
the pollen was Iteing she<l the (*ontrast between healthy and diseased catkins
wjis made mon* ronspicuous l>y the failure of infested anthers to dehisce.
SiH'tions of dise:iscMl anthers showed that, while the tissues in direct
contact with the mycelium were not killed outright, the pollen
grains were mostly degenerate, empty, and often collapsed shells. The
my(*elium is entirely intercellular, often fonning thick mats which wedge
the host cells a|)art and cause the slight <listortion of the diseased parts.
Thi>sc* myc<*lial nuits i)e(*ome especially prominent at points near \h»
surfac*(* whf*n> the large basal stroma of the fruit-body is formed in the
l(M>M* sulx'pideniuil parench>iim. Fn)m this structure the club-shaped
Ixisidia push through tlu* epidermis fonning a small but (*ompact hymen-
iiuii alxive the siu-face of the host tissue. The individual thre^ids of the
inten*ellular my(*elium and ak<o of the stnimata are extremely small and
diflirult to distinguish as »nv\\.
Ai>iMirently no toxic sulwtancc^ or injurious enz\iiies are secreted by
the fungus, siiu*e the protoplasts and nu<'lei in the infested tissue retain
nearly normal apjM'arance. The clumgi^s in cells entirely isolated by
the my<*<*lial iimts indicate starvation rather tlian toxemia. The pollen
griiiL** pn»s4»ni similar evi(h»n«*es of starvation. The vacuole gradually
enlarges an<l the* protophLxmic layer iM^^omes thinner until it disappears
entirely k'aviiig the niipty |M»llen-<*<*ll walls which collafMc or retain their
original slia|M* according to thrir degree of maturity.
IDKNTITY OF THK PAKASITE
Of llu* four HiMM'irs of Mi<*n)stn)ma mentioned in Saccardo's Syllcge
Fuf\^orum, M, album (Ik^m.) Sacc. occurs <»n leaves of oak, M. CycadiM
44 PHYTOPATHOLOaV [VoL. 7
Allusch. on leiivest of Cyaii rei-olut^i, M. unieriainuiii Paiiimel & Hiuiie on
U-avcB of CnicuM iinierictinus, uiiti M. Ju^iantiin (B<;riii([) Siicc. on k-avw*
of JuKliinx uiid Citrya. Sinrc M. Juglnndin wius uhniitlunt rarly in the
sprinK on U-uvoh of hickories, it wax iil oix-e siiHiK-ctod timt \\\c fungiin on
)K.-(-an <'atkins was iilc-ntirul wit)i tliis sjMM'ifN. Coinjiarulivc nicusurt^
nicnis of llic variiiiL" Htructurcs of the funftu;' from the two hostw Hhowctl
however Monie very markeil <ljfTerenc<'s. Tin- siMiros from [wean catkins
.-.re eyliii.lrieal. !> to M l.y .jji, an.l tlios.^ from liirkory leav.-s are oval lo
^ Pa
'o9
e CP
m°
^9
,1. rtuiOIiK l""Jv -li.miiiK l.u-i.liii. Ml.Ti|iiM:iI:i. :ii>rl -|«,r.'^: H. fr.-^li -|.i.r.-<; T.
vr».|-lil.<' r.II. rr.>ni .'itilit-.l»v»'.l.l .-iil)tir<- ..n n.rii iii.'m] -.K^af. It. <'.-IU from ..I.I
.■»Uu,.- ..I, ,.,nt iti.-:>l i.Knr. All X T::>.
ol.]iii>i!. l> U> S l.y i..V' The )ia><i<lta iw well as tlu' Klr<>ni:ila from which
thiy arjsi' :iri' innili larfcer iiiiil ihe iinniNr of lia.-<i'lia fmni each stroma
miicli icre.-iler ou ihe |M-cari.
< nlliiro i>f Uitti forms were ohiaincil arnl eom|)arei| on v:irions media.
'Ih.- r.-iK.ii-^v wen- verv similar in each ea--e. (In all nic.lia «. far Irictl.
..nh A >e:iM.|ik<- growth is formc.l. The '^porc^ -well eoi.si.lerahly an.l
wdIuii a f.'u h.inrs U-^in forminK new cells l.y a hniiiliriK jiriKi-sN, stnin
fi.tnutiK :i Mii:<1l. i-ircnlar white c.ilnnv. Ai tirsi short hvj.hal uerm hiln'<
^^.■„■ |.nnul ..<-'':.siunallv, l.ni ihcv HK.n .lis^ipixarcl, A-. th.' hii.l.linK
I.MHV^s ...nil - Ihc .taiiKliter cells lM'.'<.me smaller and ..val or clUjitieal
1917] HiGGiNs: Disease of Pecan Catkins 45
to globose. The fungus is not long-lived in cultures, requiring frequent
changes to new media.
Early in May direct and cross inoculations were tried with fresh spores
from each host, but they all resulted in failure.
Diligent search, for diseased catkins was made over several hickory
trees the leaves of which were infested with Microstroma but none were
found. Neither was any disease found on the leaves of pecan trees,
although in some instances they were almost in contact with diseased
hickory leaves.
Notwithstanding the apparent difference in tissues attacked, in size
of spores, basidia and so forth, it still seems doubtful that the pecan fungus
should be given specific rank. The more robust habit on the catkins
may be due to the more abxmdant supply of food. Therefore for the
present, it seems best to consider the fimgus on pecan catkins as merely
a robust variety of Microstroma Juglandis (Bereng.) Sacc. and to present
the following diagnosis:
Microstroma Juglandis (Bereng.) Sacc. var. robustum n. var.
Host tissue pale, often slightly distorted; myceliimi intercellular, form-
ing more or less dense mats between the host cells; fruiting stromata oval
to short conical, 60 to 100 by 55 to 150/*, compost of very slender inter-
woven threads; basidia club shaped, 13 to 30 by 5/*, bearing apically 6
to 8 spores on short sterigmata; spores hyaline, one-celled, cylindrical,
rod-shaped, 9 to 14 by 3 to 5/*.
Hab. On stamens and staminate bracts of Carya iUinoensis.
Microstroma Juglandis (Bereng.) Sacc. var. robustum n. var.
Stromaiibv^ frudificantibus subepidermids, ovatis vel brevo-conids, 60 to
100 by 55 to 85 n; basidiis caespitosis, davatis, 13 to 30 by 5^., seanf. odoporis;
sporidiis hyalinis, cylindricis, 9 to 14 by 3 to 5ti.
economic importance op the disease
Since pollen is always produced in super-abundance by pecan trees
the loss of a comparatively large amoimt is of Uttle importance. Since
however, on some trees fully one-third of the pollen was destroyed one
can readily see how the disease may become serious in the near future.
At present so Uttle is known as to the life history of species of Microstroma
that any suggestion as to control measures is almost valueless.
Observations in the Station orchard during the past spring indicated
that few or no commercial varieties are entirely immune; but the attack
was much more severe on some varieties than on others. Similar observa-
tions were also made in orchards around Albany, Georgia.
Geoboia Agricultural Experiment Station
Experiment, Georgia
SOME NEW OR LITTLE KNOWN HOSTS FOR WOOD-
DESTROYING FUNGI
Akthuk S. Rhoadb
Despite the great extent to which wood-destroying fungi have beeo
collected, but comparatively little attention has been paid to the host
species on which they occur. One frequently finds in herbaria good
collections the practical value of which is greatly reduced by being de-
ficient in this respect. Within the last few years, however, increasing
attention is being paid to the host species with the result that many
new hosts have been established and many fungi which formerly were
thought to be confined entirely to the wood of deciduous or coniferous
trees are now kno^-n to occur on both.
In his ovn\ collecting work the writer always has been particularly
interested in the hosts for wood-destroying fungi and frequently collects
for host species alone. In looking over his lists recently a few species
were noted, some of which apparently never have been reported. All
but two of the collections cited here have been made by the writer him*
self or in conjunction with others, cither in the states of Pennsylvania or
New York. The following host species for wood-destroying fungi are
believed to be new or at least little known.
Coriolus versicolor^
On dead trees, fallen trunks, and Atumpd of Tsuga canadensis* (Pa. and N. Y.).
On rustic fence rails of Juniperus vtrginiana (Pa.).
On fallen trunks and stump of Abies baUamea (N. Y.). This species was noted
as a host by Dr. L. H. Pennington on two occasions in the .\dirondack region.
Coriolus nigromarginatus
On a dpa<l trunk of Tsuga canadensis aMsocJAtcd with Coriolus abietinus (N. Y.).
Coriolus prolificans
On d(*ad trunks of Tmuja canadensis (Pa. and N. Y.). Numerous sporophores
occaAiorially are found either pure or asfl4N*iBttHl with (\yriolus aitietinus. It pro-
* The nomenclature for fungi used in this paper is that of William A. Murrill.
((Agaricales) Polyiwraccae (pars). North Am. Fl. 9: 1-7*2. 1907; (.\garicales)
Polyporareae ironcl). North Am. Kl. 9: 73-131. IIKW.I
* The nom«*nrlaturc for tr(»<*s \\mh\ in this paper xn that of George B. Sudworth.
(Check lint of \\u* forest trees* of the Tnitwl States, their names and ranges. U. 8.
Dept. Agr , Div. Forentry Hul 17: 144 p. IHIW.)
1917] Rhoads: Hosts for Fungi 47
duces a sap-rot in hemlock that is indistinguishable, macroscopically at least, from
that caused by Cabietinus,
CorioleUus sepium
On rustic fence rails of JuniperiLS virginiana (Pa.).
On stump of Tsuga canadensis (N. Y.).
Tyromyces comus
On rustic fence rails of Junxperus virginiana (Pa.).
Bjerkandera adiista
On rustic fence rails of Juniperus virginiana (Pa.).
On stump of Thuja occidenialis (Pa.) .
Porodisculxis pendiUis
On branches of fallen trunks of Juglans cinerea (N. Y.). This fungus usually is
collected on wood of Caatanea dentata but occurs commonly about Syracuse on butter-
nut wood.
PolyporiLS Polypcrus
On slash of Tsuga canadensis (Pa.).
Pycnaporus cinnabarinus
On a fallen sapling of Tsitga canadensis (Pa.).
On log of Picea rubens in corduroy road (N. Y.).
Hapalopilus gihrus
On a dead sapling of Tsuga canadensis (Pa.).
Ischnoderma fvliginosum
On dead trunk of Pinits strobus associated with Coriolus abietinus (N. Y.).
Elfvingia megaloma
On dead trees, fallen trunks, and stumps of Tsuga canadensis (Pa. and N. Y.).
On stump of Abies balsamea (N. Y.).
Ganoderma Tsugce
On or in close contact with stump of Pinus rigida (Pa.).
On stump of Picea excelsa (Pa.)-
On a much decayed stub of Betula liUea (N. Y.). A fine large specimen was col-
lected on the latter host at Cranberry Lake, New York, and was fully as typical as
those frequently found on hemlock trunks in that region.
GkeophyUum trabeum
On rustic fence rails of Juniperus virginiana (Pa.).
48 Phttopatholoqt [Vol. 7
GhBophyUum hirsutum
On A soft maple log in a wharf at Oneida Lake, New York. Occasional aporo-
pborea were anoetated with its near relative, Gloeophyllum trabeum. Tlie wood
was either that of Acer $aecharinMm or Acer rttbrum^ but judging from the dominance
of the silver maple in the lowlands of this region, it probably was the former species.
On stump and adjacent log of PrunuM avium (Pa.)* Near State College, Pa.,
•porophores were found at various times associated with Gloeophyllum irabtMm on
a stump and nearby log from the same tree.
On fallen trunk of Betula luUa (N. Y.). A collection of this plant was found in
the herbarium of the New York State College of Forestry and, although no host
was recorded, a few s|>prophoree had ample bark attached to them to be positively
certain that they grow on yellow birch.
GkBoporua e(mchoides
On an old sporophore of InonoluM dryophilue (Pa.). In making a collection of
the former plant from a black oak log an old sporophore of Inonoiui dryopkUuB waa
found on the log that also was well covered with sporophores of OUdoporuM coneAoidss
and seemed to be as good a host for this plant as the wood of the log.
The New York State College op Forestry
At Syracuse University
Syracuse, New York
NOTES ON CRONARTIUM COMPTONLE III
Perlet Spauldinq
In 1908 the writer collected a specimen of Cronartium CamptonuB
Arthur on a young tree of Pinus rigida at Burlington, Vermont. Since
that time all available information concerning this fungus has been ao-
ctunulated, and all the experimental work possible has been done. Three
brief papers^ have been published giving some of the more important
facts which have been learned. It is proposed in the present paper to
state very briefly some results secured in more recent investigations.
The pine hosts of Cronartium Comptanias, which have been previously
reported by various writers, are Pinus rigida Mill., P. sylvestria L., P.
maritima R. Br., P. austriaca Hoess., P. divaricata Ait., P. echinata Mill.,
P. montana Du Roi, P. ponderosa Laws., P. contorta Loud., P. virginiana
Mill., and P. tceda L. In the years 1915 and 1916 Cronartium Comp^
tonicB has been received by the writer from various locaUties on the follow-
ing new pine hosts: Pinus densiflora Sieb. & Zucc. (one locality), P.
jeffreyi Oreg. Comm. (two localities), P. larido Poir. (one locahty), P.
mugho Poir. (three localities) and P. resinosa Alton (one locaUty; three
other localities are known but no specimens could be secured).
Successful inoculations have been made by the writer and his colleagues
named below, for the first time, so far as can be determined from pub-
lished statements, from several pines to the alternate hosts. Uredinia
were produced on plants of Comptonia asplenifolia L. with aeciospores
from Pinus tceda (one test made), P. austriaca (2 tests), P. rigida (4 tests)
P. mugho (3 tests), P. resinosa (1 test), P. jeffreyi (1 test), P. larido (1
test). Uredinia on Comptonia were successfully used to produce uredinia
on Comptonia and Myrica gale, Uredinia from Myrica gale produced
uredinia on Comptonia. These are the first successful inoculations with
uredinospores to be reported. In the winter of 1914-15 a special effort
was made to seciu'e Uving plants of all the species of Myrica growing in
this coimtry. A stock of the following species was obtained largely
through the efforts of G. G. Hedgcock and E. P. Meinecke: Myrica
gale L., M. califomica Cham., M, cerifera L., M. carolinensis Mill., M.
'Spaulding, Perley. Notes on Cronartium Comptonise. Phy topath. 8 : 62. F. 1913.
. Notes on Cronartium Comptonise II. Phy topath. 8: 306-310. D. 1913.
. Notes on Cronartium Comptonise and C. ribicola. Phy topath. 4: 409. D.
1914.
50 Phytopathology (Vol. 7
inodora Bartr., M. pumila Michx., as well as CompUmia asplenifolia.
Mr. G. F. Gravatt and Dr. G. R. Lyman in 1915 made inoculations under
the writer's direction. Five different series of inoculations, with aecio-
spores from as many different species of pine, were made. Theresulta
were as follows:
3 plants — CompUmia asplenifdia — uredinia produced on all.
7 plants — Mj/rica gale — ^uredinia produced on all.
9 plants — Myrica caroHnerms — no infection.
4 plants — Myrica califamica — no infection.
5 plants — Myrica inodora — no infection.
3 plants — Myrica pumila — no infection.
4 plants — Myrica cerifera — no infection.
A single plant of 3/. carolinensis was inoculated repeatedly in 1916 by
the writer without visible results other than yellow spots on the leaves,
which are thought to be due to some other cause. Nimierous successful
inoculations have l)een made on CompUmia asplenifoUa: i.e., a total of two
in 1912, eleven in 1913, two in 1914, nine in 1915, and fourteen in 1916.
A lesser numljer has been made on Myrica gafe, which gives much less
striking results. The total number of such successful inoculations on
Myrica gale is: seven in 1915 and four in 1916.
In 1912 the writer l)egan a series of annual observations in an area of
several acres near I^kc George, New York, quite thickly covered with
natural reprcKluction of Pinus rigida, in which Cronariium CompUmia
occurs as a native parasite. The size of the trees within the area ranges
from twenty feet in height downward. Ver>' few are as small as two feet
in height and there seem to l)e no ver\' young see<llings now appearing.
Comptonia ojsplenifolia gro^-s naturally throughout the area so that condi-
tions are excellent for the spread and development of the disease. These
observations have continue<l until the present time, thus covering a period
of five years. An effort has been made to keep numl)ered labels on all of
the (iis(»jised trees, hut with indifferent success Ix^cause of curiosity or
mischief in people who luippened to see them. Several significant facts
liave l)een learned howevcT. On pines Cronartium CompUtnict fruits for
a iK»ri(Ml of seven or eight w<M»k.**, the time of maximimi fruiting being
alMMit Jiine first <'on8iderably earlier than the Tiriter at first supposed.
The numlK»r of diseasc^d trcH»s killed annually by the fungus has been sur-
prisingly unifonn, mnging from eight to eleven. In 1916 the obecrva-
tion.H were made earlier than iLsual and a greater total niunber of diseased
inN-s were unexi>e<'t<»<||y found. There is evidence that a small munber
of tr(H»s are annually infeeted for the first time and it is hoped to learn
this numtxT in the future. It is well establi.Hhed tliat a tree which once
1917] Spaulding: Cronabtium Comptoniae 51
bears fruit of the fungus almost always bears an annual crop of such fruits
until the tree dies. In some cases a dying tree fails to produce them,
but on the other hand, recently killed trees are often found with aecia
on them. While the annual loss is not great, it is a serious loss when
continued indefinitely. Moreover we have excellent reasons for believing
that Pinvs rigida, to which the preceding statements apply, is much less
susceptible to the disease than are P.^onderosa and P. cantarta. The loss
in some instances, with the latter two species, has been total.'
Ofpicb op Investigations in Forest Pathology
Bureau op Plant Industry
Washington, D. C.
'Kauffman, C. H. and Mains, E. B. An epidemic of Cronartium Comptonis at
the Roscommon state nurseries. Mich. Acad. Sci., Ann. Rept. 17: 188-189. 1915
FURTHER NOTE ON A PARASITIC SACCHAROMYCETE
OF THE TOMATO
Albert Schneider
Since the appearance of the recent article on a parasitic Saccharomyoete
of tomato^ it has been determined that the fungus described unquestion-
ably belongs to the genus Nematospora of Peglion.' In its morpho-
logical characteristics it is closely similar to Nematospora Coryli PegL,
which attacks the fruit of the hazel bush (Carylus aveUana). Peglion,
however, makes no reference to the gametic origin of the ascus nor does
he note the two cells of the ascospore. He also fails to recognise the
arthroHpores and includes them imder ''Anomale vegetative Formen."
These and other morphological as well as biological differences make it
clear that the Xematospora Coryli of Peglion and the fungus under con-
sideration are two distinct species. Both are true parasites and appear
to occiu- in wanner countries (southern Italy, southern California and
Cuba) and perhaps also in semitropical and tropical countries.
The Nematospora of the tomato (Lycopersicum esculentum) is appar-
ently a new species and the following name is therefore proposed:
Nematospora Lycopersid n. sp.
Asci of gametic origin soon becoming free from associated cells, cylin-
drical with rounded ends, 60 to 70m in length; ascospores in two groups
of four spores each, two-celled, slender, with pointed ends, slightly ridged
at transverse septum, 50 by 4.5^; ascospores liberated by dissolution of
ascus wall and held together somewhat in groups of 4 by motionless fla-
gellae; flagellae 50 to lOO/i in length; arthrospores, of non-gametic origin,
spherical to ampulliform, 25m in diameter. Two other cell forms also
found: (1) much elongated, filamentous cells; (2) elliptical and ovoid cells,
gametic in function, new celU forme<l in bipolar direction by apical bud-
ding and also by apico-lateral budding at cell unions. The elliptical and
ovoid cells alone are gametic in function.
Habitat. Parasitic on nearly ripe and ripe fruit of Lycopersicum es*
culenium, southern California, Cul>a and Mexico.
'Srhnridcr. Alljert. A paranitic Saccharomycetc of the tomato. Phytopatholofj
4: 395-399. 1910.
•IVglion, Viiiorio. ToJior dir Ncmatoapora Coryli Pegl. Ccntralbl. f. Bakt.
Abt. 2, 8: 754-761. 1901.
1917] Schneider: Parasitic SAccHAROBiTCETE of Tomato 53
Nematospora Lycopersid sp. nov.
Ascis cylindratis, terminate orbiculato; 60-70/i in longitudine; sporidiis
8; dispositis struibus duo, sporidiis 4. Ascus mox ex cellis prehensis li-
beratus est. Tnnicae asconun in maturitate solventur et sporidiis liber-
antur. Plurimi asci origine gametata sunt.
Sporis ascpnim bi-cellulatis tenuibns, fusiformibus, flagello uno. Spo-
ridiis 50 X 4.5m, flagello 50-100m.
Arthrosporis non-gametatis, plerumque sphaeroidis, 25/i.
Cellulis vegetativiSy eUipticis, ovatis ad filaris nonramosis, multipli-
cantibus gemmatando apiculo-laterali apicnlatoque. Cellulis ellipticis et
ovatis solis sunt gametatis.
Hab. In fructo Lycopersici esculenti, terris calidis et tropieis.
Acknowledgments are hereby made to Dr. Roland Thaxter and Prof. H.
W. Anderson for calling the writer's attention to the Nematospora of
Peglion.
Uni\ersity op California
College of PnARBiACY
San Francisco, Calif.
PHYTOPATHOLOGICAL NOTES
Albany conference on white pine blister rust. A joint meeting of the
North American Committee for the Control of the Pine Blister Rust and
the cooperators of the United States Department of Agriculture was held
at Albany, New York, November 20 and 21. The meeting was attended
by a representative body of men from the states of New Hampshire,
Vermont, Massachusetts, Rhode Island, Connecticut, New York, New
Jersey, Pennsylvania, Virginia, Indiana, Wisconsin and Minnesota, and
from the Dominion of Canada and the United States Department of
Agriculture. The state foresters, pathologists, and nursery inspectors most
intereste<l in the fight for the control of the disease were present.
A brief re|x>rt from each state and from Canada gave the latest news
concerning disea^ survey and eradication work. The reports show the
general epiphytotic of the blister rust in New England and eadt of the
Hudson River, with comparatively few centers of infection in New York,
Pennsylvania and New Jersey, one infection center in Ohio, two in Wis-
consin, and four in Minne«K>ta. A hasty survey made of the western
half of the count r>' faile<l to show any blister rust of white pine present.
It developed tliat the matter of control of the blister rust in the New
England states resolved itself into the practicabilit>' of the eradication
of currant and gooseberry' bushes on a large scale. The control of the
blister rust in the I^ke States was shown to be on a somewhat different
basis, for in that region the spots of infection known are very few. Here
the total eradi(*ation of all white pines and Ril)e8 near the infection cen-
ter is l>eing carried out.
The Conmiittee passe<l resolutions favoring adequate legislation which
would |)ennit states to carr>' out the eradication or control work neces-
sary', following largely the Sanders model horticultural inspection bill.
They also favored ad«|uate appropriiitions by the states to carry on
eradication work and by the Fcnleral (Sovernment to carry on the sur-
vey an<l ex|KTimental work. The Committee favored a Federal quaran-
tine prohibiting shipment of five-needled pines and all species of Ribes
from the eastern half of the countrj' to any part of the country west of
Minm^sota, Iowa. Muwouri. Arkansas and Ix)uiBiana. A state qxiar-
antine was also reconanended pn>hibiting shipment of five-needled pines
and RilK»s from inf(»<'ted stat<»s to others not infected.
A national law was urfccnl prohibiting the importation of all plants
1917] Fhttopatholoqical Notes 55
from any other continent into the United States except through the
United States Department of Agriculture solely for scientific and experi-
mental purposes.
Rot G, Pierce
Corn disease caused by PkyUachora graminis. During the euminer of
1915, an apparently undescribed disease of corn was observed by F. L.
Stevens in Porto Rico, Leaves were collected from the diseased fields
in numerous localities in Porto Rico, and a study of the disease based
entirely on this herbarium material has been made at the University of
Illinois.
Fig. 1. Pbtllacrora graiunis on Corn
PortioD of a leaf shows strcmata on upper and lower surfaces
The disease manifests itself as well-defined, aubcarbonaceous spots,
either dmall and numerous or comparatively large and sparaely distributed,
and in either cade surrounded by a narrow, yellowish-brown halo. The
infection is local, confined to the leaf and the leaf sheaths, and is visible
on both upper and lower surfaces.
The spots are due to the formation of stromata in the infected tissue.
Embedded in the stromata are perithecia consisting of depressed cavities
surrounded by walls made up of dark-brown mycehum and bearing at
the top a comparatively small ostiole.
The mycehum in newly invaded tissue is slender and hyaline. It
later becomes dark brown, filling the leaf tissue of the infected area with
a network of hyphae.
66 Phytopathology [Vol. 7
The asci are numerous. They are cylindrical and in each sBcua are
produced eight unicellular, hyaliqe, thin-walled spores.
The fungus causing the disease has been identified as PhyUaehara
graminus,
A report including a detailed description of the disease and fungus,
together with notes concerning the generic and specific relationship, will
be read before the Illinois Academy of Science at its next meeting.
Nora E. Dalbey
Tylenchus triiici on wheal. In August, 1915, specimens of diseased
wheat plants, which had just been received from Dr. J. H. Reisner, of
the University of Nanking, Nanking, China, were kindly turned over
to the \vTiter by Dr. H. B. Himiphrey, of the OflSce of Cereal Investigations,
Bureau of Plant Industry. In transmitting this material by letter Dr.
Rei^ner said, *'Tlie disease has become more widespread every year for
the last three or four years and is causing great money losses."
A microscopic examination of the wheat heads showed that practi-
cally all of their glumes contained, in place of normal kernels, dark galls
filled with an almost innimierable number of motionless but living larvae
of the nematode, Tylenchus triiici Bauer, which has been known as a serious
pest in Kurope since 1745. The parasite has been found in Sweden,
Holland, (lemiany, Austria-Hungary, Switzerland, Italy, England, and
Australia, but so far as known has never been reported from China before.
Johnson* recorded in 1909 the occurrence of what undoubtedly is the
same species on wheat from a few widely separated sections in the United
States, but as no reports of its appearance l>efore or since that date have
iK'on found, it is quite unlikely that the eelworm has become well estab-
HsImmI in the wheat areas of this country. Whether the closely related
siH»ci(»s of Tylenchus found by Bessey*, the writer, and others on several
difTcrent grasses in various parts of the United States is identical with the
fonn on wheat has not been determined. Some European investigators,
however, regard Tylenchus triiici as a highly specialized parasite of the
wheat.
The infected heads of wheat are usually shorter and thicker than nor-
mal heads and contain glumes which spread out almost at right angles
to the fruit stem. In place of normal seed, dark, hard galls, incapable
of g(*rmination ami full of lar\'ae, are to \ie found. Because of these
effects on the host the disease has merited such descriptive names as ear-
coc'kles, purples, false ergot, etc.
* Johnwm. VAw, C NuU'S on a nomatodo in wheat. Science n. a. SO: 576. 1900.
* B^aaey, Krneat A. A m*matodc* diaeaap of graasea. Science n. a. 21 : 391. 1006«
1917] Phytopathological Notes 57
Active larvae enter the young, tender tissues of the wheat flower,
extract -food therefrom, tnature, and lay eggs, which in tiun give rise to
another generation of larvae. After reaching a certain stage of develop-
ment some of the subsequent generations of larvae become coiled and
dried out in the matured seed coats of the host and are capable of re-
maining in this inactive condition for long periods. Under favorable
conditions of moisture and temperature the eelworms may escape from
the seed, attack the leaf and stem parts of wheat seedlings, causing them
to become wrinkled, distorted, or swollen, and finally enter the embryonic
seeds.
It has seemed desirable to bring the above data to general attention,
in the hope that active measures will be taken both to prevent the in-
troduction of this parasite along with wheat importations from infected
coimtries and to stamp out the pest wherever it is foimd in this coimtry.
L. P. Byars
The Botanical Society of Washington. The following officers have been
elected for the ensuing year:
President, Mr. T. H. Kearney; Vice-President, Mr. Edgar L. Brown;
Recording Secretary, Mr. Charles E. Chambliss; Corresponding Secre-
tary, Dr. H. L. Shantz; Treasurer, Mr. F. D. Farrell.
Mr. A. S. Hitchcock was nominated by the Society for the position
of Vice-President of the Washington Academy of Sciences.
Personals. Mr. Chas. S. Reddy, of the University of Wisconsin, has
been appointed as assistant plant pathologist, and Mr. A. M. Christen-
sen, of the North Dakota Experiment Station, as an agent, in Cereal
Disease Investigations, Bureau of Plant Industry, with headquarters
at Fargo, North Dakota, where they are engaged in the investigation
of cereal diseases in cooperation with the North Dakota Station.
Mr. F. A. McLaughlin, instructor in botany at the Massachusetts
Agricultural College, has been granted a year's leave of absence for grad-
uate study at the University of Chicago.
Mr. W. L. Doran, for the last two years graduate assistant in botany
at the Massachusetts Agricultural College, has been appointed instructor
in botany and assistant botanist at the New Hampshire Agricultural
College and Experiment Station.
ABSTRACTS OF PAPERS PRESENTED AT THE EIGHTH AN-
NUAL MEETING OF THE AMERICAN PHYTOPATHOLOGICAL
SOCIETY, NEW YORK CITY, DECEMBER 26^, 1916
Evidence oj the over wintering of Cronartium ribicola. Perlet SpaulDING
Numcrouii inHtances have been noted whore largo lots of black currants were very
heavily infected with Cronartium ribicola one summer and not the next. In the
Geneva New York case the disease was present upon pines and these in the writer's
opinion started the disease each spring. Cooperative experiments with Stewart,
in which 500 heavily infected black currants were used, resulted in no disease.
Furthermore, the writer has had during the past sev(*n years in the greenhouses at
Washington, hundreds of KilK»s plants of more than thirty species, which have l>een
UM>d in inoculation ox|>orimonts. In no case has the disease ever appeared upon
thest* the next w^ason until artificial inoculations had l>oon made. Uniform success
has l)e<*n obtained |]i*rs<>n»lly in locating the diseased pines from the areas occupied
by the first gi'noration of un'doKporoM in the field. Field observations seem to indi-
cate that the dis(*am* has lieen shipfied into now localities on infected Ril)es. In-
fections of petioles are not as rare as at first supposed. No evidence of bud infection
by way of the petiole has yet Ix^en s<*cured. Direct examination of buds on infected
plants has also failod to show the presence of the disease.
The control of white piri£ hlinter rwtt in small areas. W. H. Ra.vkin
The control of white pine blister rust has l>een att4>mpted in eighty-five forest
plant ingH mad<» with imported stock in New York State. Diseased or suspicious
trees and all currants and gooseberries within five hundred feet of the plantings
were n^nowd. Thirty-six plantings ha\'e shown diseased trees since 1909; twenty
since HUl; sewnti^en sinci* 1912; fifteen since 1913; nine since 1914 and foursince
1915. The HifH-9 sp. within one-half mile of all the plantings were inspected in the
autumn of 1910. Dim>ased HH>en sp. were foumi around two, only. These were two
of the four which had shown diseased trees in the spring. In both cases cultivated
varieties of KiheM still existed within five hundred feet. It seems, therefore, that
th«' n-moval of diseased trec*s and all currants and gooselK^rrios within five hundred
fe«*t of the plantings has prc\Tnted the establishment of Cronartium ribicola in
these an*as.
Citrwt canker invcntigationn at the Florida Tropical lAiboratory, R. A. Jbhlb
S<}rne cultural charuct'*ristirtt of the canker organism, Pseudamonas Citri Haase
(Migula'rt genuH) or liactrrium (*itri (Crohn's genus Bacterium as emended by Smith)
are as follows: (trowth f>n standard agar abundant, spreading, raised, smooth,
glistening, transluscent, pale >-ellow, viscid, with characteristic odor in about
6%^ days. Vitality ten to thirty day's, (trowth on |M>tato agar more abundant
and spn*adtng with lighter color. Growth on potato slices more viscid and brighter
>tIIow with fiifltinrt white margin on |K)tato adjaoi*nt to culture. White margin
l»ecomes wry pn>mim*nt in forty-eight hours and do«>s not stain with iodine. On
gra|M*fruit leaf and st4*ni decoction agar slants grom'th |Nrnet rates the agar and if
1917] New York Meeting 59
less raised. On oat agar slants growth is much more spreading and the color is much
lighter.
Positive results have been obtained from inoculations on grapefruit, ponderosa
lemon, key lime, Citrus trifolicUOf sour orange, tangelo, sweet orange, tangerine,
king orange, mandarin-lime, and kumquat. Disease also occurs on navel orange,
mandarin, satsuma, conmion lemon, rough lemon, and Aegle glutinoaa.
Studies upon the anthracnose of the onion. J. C. Walker
A morphological study of the causal organism Colletotrichum circinans (Berk.)
Vogl. shows the fruiting body to be an acervulus and not a pycnidium as first de-
scribed by Berkeley, confirming the findings of Voglino who transferred the fungus
from the genus Vermicularia to Colletotrichum. Further study has shown it to
conform closely to the description of Colletotrichum fructum (S. & H.) Sacc. {Volutella
fructi S. & H.), reported by Stevens and Hall as causing a fruit rot of apple. Inocu-
lation of the fungus from onion into apple fruits resulted in a rot very similar to
Volutella rot. Further study is necessary before the two fungi can be considered as
identical.
Inoculation of onion bulbs in soil held at different temperatures shows best in-
fection to take place between 24** and 29**C. This may account in part for the rather
sparing appearance of the disease until shortly before harvest. The fungus over
winters in the soil and consequently the disease is most severe on old onion fields.
Spraying the bulbs before harvest or in the crates after harvest has not proved
beneficial. The fact that yellow and red varieties of onion are highly resistant
offers some encouragement for the development of a resistant white strain. Work
in this direction is to be continued.
Pink root, a new root disease of onions in Texas. J. J. Taubenhaus and A. D. Johnson
A new disease known as pink root is causing serious losses to onion growers of
Webb County, Texas. The trouble seems to prevail only where onions are grown
for two 3rears or longer on the same land. The same is also true for the seed bed
where the same old soil is used for several 3rears in succession. The disease starts
with the young sets in the seed bed and from there is carried to the field.
The roots of the affected sets in the seed bed or the plants in the field turn pink
in color then strivel and die. As fast as new roots are formed they become infected,
turn pink and dry. The effect of pink root is to prevent the normal development
of the bulbs in the field and to produce dwarfed undersized bulbs which are absolutely
worthless as far as the market is concerned. The cause of the disease is still proble-
matic. Attention is called to it at this time because of its great economic impor-
tance to Texas onion growers. Extensive investigations are now under way to
determine the cause and possible remedies for this disease.
Two new camphor diseases in Texas. J. J. Taudenhaus
Two new or little known diseases seem to threaten the existance of camphor
trees in Texas :
1. Anthracnose. The fungus attacks and kills the leaves and branches. Affected
trees have a defoliated appearance at the top. The cause of the trouble is apparently
a new species of Glceosporium tentatively named Olasosporium camphorcB. The
organism is readily grown in pure culture and the disease reproduced at will.
2. Limb canker. This disease is characterized by a dying of the larger limbs to
about four to six fe^t from the top. The limbs turn dark and soon shed their leaves.
Affected trees have a ragged and burned appearance . A fungus of the genus Diplodia
60 Phytopathology [Vol. 7
is &lwa3r8 aasociated with this disease. Investigations are now under way to de-
termine whether this Diplodia is the same or similar to D, Campkorm F. Taaai,
occurring in Italy, and whether also it is the direct cause of the disease. A full
description of the two organisms will appear at a later date.
Common and Bcientifie name* of plant dUeaiti, M. B. Waits
Common names of plant diseases are used by a larger number of people than
scientific names. Pathologists should encourage the movement to make oommon
names definite and national. By being made definite they can attain their proper
status in discussions, literature, dictionaries, quarantine regulations, laws, and
legal proceedings. By agreement among pathologists they may even become more
fixed than scientific names.
There may be four distinct names connected with every parasitic disease; the
oommon name of the disease, the scientific name of the disease, the common name
of the parasite, the scientific name of the parasite. For example: lemon scab.
Verrucosis, lemon scab fungus, Cladosporiwn Cilrx. In case of all common diseaeee
the aim should be to provide these four names. Confusion has resulted in the fail-
ure to recognise these four kinds of names, particularly in the use of the scientific
name of the disease and the scientific name of the fungus as the common name of
the disease. Scientific names may become conmion names through use but these
cases should be clearly recognised as such and avoided if possible and vacancies in
names also recognised.
Nonparasitic diseases may have two names, common and scientific, and the same
principles apply.
Economic HomU of Sclerotinia libertiana in tidewater Virginia, J. A. McClintock
The warm, humid climate of tidewater Virginia is especially favorable to the
development of ScUrotinia libertiana. This fungus, long known as a serious parasite
on lettuce, has been observed to destroy over fifty per cent of the autumn lettuce
crop on farms where no rotation is used. In the fall of 1915 a serious disease of snap
beans due to this organism was found. During the winter of 1915-1916 Sclerotinia
libertiana was found to be the cause of a fruit rot of tomato in the greenhouse. In
winter-grown parsley, under sashes, this fungus in one case caused drop of ten per
cent of the crop in the infected frames. Sclerotinia libertiana caused the damping
off of a large proportion of the seedling plants in several cold frames of cauliflower
being raised for a spring crop of 1916. In the summer of 1916 this fungus caused a
stem blight of liearing egg plants, on several farms. In each case the writer was
able to isolate the causal organism and to reproduce the disease in the respective
hosts, and in other hosts by cross inoculation.
Lima Itran monaic. J. A. McClintock
During the summer of 1916 while conducting experiments with nine varieties of
pole, and s(*\'en varietieif of buith lima Iwans, the writer observed a serious mosaie.
It wait olMier\'ed first on the Sieva pole lima or butter-l)ean and later on Improved
Hendenwm'ii Biuih and Prolific bush, lima lieans of the Sieva type.
Owr twenty-five per cf»nt of the several hundred plants of each of the above-
mentioned varieties wen* stunted and l>ore the dwarfed, mottled, wavy leaves, ehar^
actertstic of this mosaic. None of the varieties of larger limas, which made up the
remainder of the planting. showiMl signs of this mosaic, though they were grown under
the same conditions and in many cases intertwined with the mosaic diseased plants
1917] Nbw York Meeting 61
of the Sieva type. Lima beans had not been grown on this land previously and no
beans of this type were Rowing nearby, therefore, it was concluded that this lima
bean mosaic was carried by the seed. This disease is serious because the yield on
the infected plants is greatly decreased and the pods are smaller and malformed.
Bean mosaic. V. B. Stewart and Donald Reddick
Hundreds of acres of pea beans (Phaseolus vulgaris) in New York showed the mosaic
disease in 1916 and in some fields practically every plant was afifected. Affected
plants rarely set pods. The disease is not confined to pea beans. Numerous other
varieties of dry and snap beans showed the disease but not so commonly as pea beans.
The mosaic-diseased leaves on affected bean plants show irregular crinkled areas,
somewhat deeper green in color than the surrounding 3rellowish green tissue. The
disease is transmitted through the seed. Bean seed from mosaic-diseased plants
developed diseased seedlings. Healthy seedlings rubbed with crushed mosaic-dis-
eased leaves showed infection four weeks later. The first signs of the disease ap-
peared in leaves which developed about blossoming time. Leaves which had de-
veloped previously remained healthy. High temperature |knd humidity at time of
inoculation slightly favor infection.
Two transmissible mosaic diseases of ctxumbers, Ivan C. Jagger
In Phytopathology for April, 1916, there is a group of articles, dealing with a
mosaic disease of cucumbers, commonly known as white pickle, which causes a mot-
ling of both leaves and fruits. In the vicinity of Rochester, New York, there occurs
a second and distinct mosaic disease, which exhibits a mottling of the leaves, but
shows no effect on the fruits. The latter disease has been repeatedly transmitted to
healthy plants by rubbing with crushed diseased leaves, and has been transmitted
to muskmelons and to summer-crookneck squashes. This may be the disease
observed by Selby in Ohio and by Stone in Massachusetts.
Bean diseases in New York State in 1916. W. H. Burkholder
An investigation of the diseases of the field bean in western New York begun in
1915 was continued during the summer of 1916. The most serious disease, a dry
root rot, caused by a species of Fusarium, was reported last year. Morphologically
the pathogene is nearly identical with Fusarium Martii Ap. and Wr., although
infection was not obtained by inoculation with the latter fungus. The organism
winters over in manure where bean straw has been used as feed, and thus may be
spread from field to field. There is also evidence that the fungus may live for many
years in the soil. The disease was foimd in practically all of the one hundred and
fifty fields visited in western New York. Apparently all varieties of beans are
equally susceptible to the disease although certain undesirable types of the white
marrow are very resistant. A few individuals of these have been selected for breed-
ing stock.
The blight, caused by Bacterium Phaseoliy and the mosaic along with dry weather
also aided in reducing the bean crop of 1916. There is some indication that Bac-
terium Phaseoli causes a stem girdling. Anthracnose was destructive in 1915, but
caused little damage in 1916.
Do the bacteria oj angular leaf spot of cucumber overwinter on the seed? Eubanks
Carsner
This question was first suggested by the writer's observation in June, 1915, of
the occurrence of angular leaf spot in a field on recently cleared land surrounded
6i Phytopathology [Vol, T
bj woods near Portsmouth, Virginia. This field was removed at least three or four
miles from any other cucumber patch.
In 1916 near Madison, Wisconsin, six separate experimental fields were planted
with seed from the same source on land which had not been planted to cucumbers
for at least three years. The disease appeared on seedlings in all of these fields and •
in three of them it was noted on the cotyledons. In three commercial fields in the
same vicinity, planted with seed from other sources, the disease did not appear
at all in one case and not until late in the season in the other two.
The fart that angular leaf spot appeared on seedlings only in the six fields planted
with seed from the one source and not in the other fields in the vicinity furnishes
the basis for the working hypothesis that the causal organism is seed-borne, and is
oppo8C<i to the theory of local overwintering of the organism by means of insects or
plant debris.
Infrrte<l cucumber fruits in considerable numbers have been seen by the writer
in seed fields. The method of securing and cleaning the seed affords ample oppor-
tunity for the organisms to reach the see<l, and the process includes no operation
that would be likely to |^ill all of the bacteria.
Prelimiuary notcn on n ttetr leaf np<tt of cucumhern. (Iko. A. Osnkr
During the 8oa.'<ons of 1915 and 1910. the writer's attention was called to a peculiar
leaf spot on ruruinbcrH that was raiising more or leH.*< damage in a number of fields.
The sp<»t8 variwi from two tenths to fiftt»en millimeters in diameter, the majority
of the smaller spots ranging from on<' to two millimeterH and being limited in most
cases by the veins of the leaf. The larger spots were white or tingcxi with brown
and with rcnldish brown areas along th<' veins of the leaf which gave the sjMits a
chararteristir mottleil appearance.
The dinea^e was found to be due to a fungus belonging to the DematiaceavDictyo-
spone group of the Hyphomyretes. Its exact generic position has not been deter-
mineti a.** yet. The <irganisni was securcnl in pure culture on string bean agar and
8uc<'e*«sful inorulntions have been made on young cucumber plants in the green-
house, the rherkM remaining healthy in all eases.
ViruUncf of Hiffrrrut ntraiuH of CUuiosporium ciwumrrinum. W. W. GiLBEKT
\ considerable number of strains of the cucumber scab fungus, Cladonporium cm-
cumrrinutn. have been isolatiNl from cucumber fruits from different localities and
grown in culture, and many series of inoculation experiments have IxH'n mmle tm
sei*<llings and young and old plants. It has been found that these strains vary
widely in their ability to infect cucumber plants. Some have failtnl to prcnluce
any infertion after many trials, while others are uniformly virulent. One strain
in partifiilar liaj* ^iven very striking infection practirally I'very time when favorable
conditionM were maintain***!. The virulent strains attack cotyle<lons and stems of
young cucumber ^e<Nllings in inoi.st chamlxTs and kill them in two to four days.
They also attack and kill vtTy rapidly the younger leav(*s, stems and growing tips
of larger iilants under favorable moisture conditions. I'nder identical conditions
the iionvirulent strains faihsi to prcnluce infection. Similar results were secured
when youiiK cucunib<>r fruits were inoculated in moist cluunbers with different strain*
of (*lado^|M>rium.
Ihfntmxuotiitu ♦»/ Ihr nrgnninm of rurumher anlhracnone . M. \V. GARDNER
In A fairly large acreage of cucumbers and other cucurbits under observation
in 19HV the ap|M*arance of original centers of anthracnose in only the plots or fields
1917] New York Meeting 63
planted with seed from certain sources pointed suspiciously toward disease intro-
duction with the seed. Subsequent inspection of seed farms revealed the presence
of anthracnose on seed fruits and that the processes involved in seed extraction may
furnish efifective means of seed contamination.
Extensive spread from original centers of infection in the cucumber fields under
observation followed periods of heavy rainfall. The principal direction of spread
was that of the surface drainage. Plate isolations of the fungus were made from soil
near diseased plants. Rows of healthy seedlings exposed to drainage water in dis-
eased fields during heavy rains the first week of September became abundantly
infected and many seedlings were killed outright.
Glass tumblers were sunk in the ground at various points in two fields to intercept
surface drainage during rains. A successful plate isolation of the fungus was made
from water thus collected. Samples collected in one field after rains on Septem-
ber 5 and in the other field after rains on September 12, when sprayed or sprinkled
on healthy potted cucumber plants, produced numerous anthracnose lesions.
A bacterial stem and leaf disease of lettuce. Nellie A. Brown
A serious stem and leaf disease of lettuce appeared in Beaufort County, South
Carolina, 1916. The disease occurred chiefly on two plantations, one of seven-
teen, the other of nine acres; a conservative estimate of loss on the former was sixty
per cent; on the latter ninety per cent. Other plantations within a radius of twenty
miles suffered one to ten per cent loss.
The affected plants were wilted, the outer leaves blotched and darkened. Rot-
ting was often rapid. A cross-section of stem at an early stage of disease showed
a blue-green color; later stage, brown. Both pith and vascular regions were in-
volved. Later the stem usually became dry and brittle. Moderately diseased
plants are darkened in patches in stem, and vascular region. Bacteria filled the
cells of the blue-green and brown areas. No fungi were found.
A bacterial organism was isolated which when inoculated into lettuce produced
the blue-green color throughout the vascular system and pith, which later became
brown. Disease appeared on leaves also. Eight months after isolation this or-
ganism, which is yellow on various media and is doubtfully motile, is still infectious.
In its morphological and cultural characters it does not correspond with any or-
ganism recorded as pathogenic to lettuce.
Blajck spot of pepper. L. E. Melchers and E. E. Dale
In 1915 a striking pathological condition of fruits of peppers was noticed; the
disease ranging as high as 45 per cent. A species of Altemaria has been consistently
associated with diseased areas. These are slightly sunken, dark colored and not
confined to any particular location. Inoculations show that the organism is only
a weak pathogene when inoculated into normal tissue. When the pericarp is me-
chanically (slightly) injured, the fungus becomes established and diseased areas
enlarge. There are apparently two ways or combinations of factors in which natural
injuries may occur to peppers in the field; (1) injured areas from sun-scald, (2) frost
injuries. Artificially injuring the pericarp by means of a burning-glass and apply-
ing the fungus superficially, produces a condition which is the counterpart of the
symptoms occurring in the field. Varietal resistance is shown by the data of 1916.
Sweet peppers are more susceptible than the hot varieties. Among 15 varieties
grown, the percentage of disease ran from 0.02 to 2.7 among hot peppers and from
0.4 to 23.07 among the sweet ones, with an average of 13.6 among the latter when
sprayed and 11.7 in the unsprayed. Bordeaux sprays do not control the malady.
Affected fruit is unsalable.
64 Phytopathology [Vol. 7
A 9cUrotium dUeoMe of pepper$. Wiluam H. Martin
The disease is characterised by the presence of numerous, minute, blaek sclerotia
throu|(hout the fruit as well as on the seeds. With the exception of a blackening
of the epidermis, the disease may pervade the entire interior before any external
symptoms are noticed.
The fungus was isolated and grown in pure culture. The pathogenicity of the
fungus has been fully established by numerous successful inoculations on both
green and ripe fruit, as well as on the plant itself, and by the subsequent re-isola-
tion in pure culture. Reinoculations with this second isolation again produced
the typical rot.
The identity of the pathogene has not been satisfactorily determined but there
is evidence for the belief that it is ScUrotium bataticola Taubenhaus.
Successful cross inoculations have been made on pepper and sweet potato as well
as on cucumber, tomato, apple and egg plant.
DiMfffiifui/t on of Bacterium hfalvacearum, R. C. Faulwetter
Through investigations of the means by which Bacterium hfalpacearum may be
disseminated, it has been concluded that wind during rainfall is the most impor-
tant agent. Neither insects nor seed infection can account for the prevalance of
the angular leaf spot caused by it in all varieties of cotton. An inoculation experi-
ment consisting of one row of plants in a field free of the disease was followed within
a month by infection of the plants to the east as far as the fourteenth row, and in
the next month by the spread of the disease to the west. A second experiment ar-
ranged and operating during the second month also showed spread to the west.
Practically no disease occurred east or west of uninfected plants in the original
rows.
It was demonstrated experimentally that the leaf-surface film during heavy dews
contained viable bacteria. It is to l>e expected these organisms will be present
during rains. Westerly winds prevailed during the rains at the time the disease
spread to the east, and during the next month the wind direction had changed, blow-
ing from the south-east. Considering the slight disease opposite the uninoculated
plants, and the simultaneous change of wind direction and the spread of the dis-
ease, it is held that wind during rainfall is the most active agent in the dissemina-
tion of the causal organism.
•
Bacterial dineaseM of celery. W. 8. Kroitt
These diseases occur in a region with a deep muck soil and a very humid climate.
Trotrn rot. This disease is prevalent in the greenhouses and fields. The symptoms
are a yellowing of the foliage and a rotting of the crown starting through side roots.
Plants arc destroye<l in fnim three to four wcekn after infection. The causal organ-
isms appear to l>e a Bacterium and a Fui«ariiim working simultaneously. Steam
steriliiation and the following rhemimls have lieen applied in varying amounta to
the soil for the control of the disease: foniialin, calcium chloride, copper sulphate,
ferrous sulphate, sulphur, scMliuni chloride and potash.
Crown rot trilt. This diHoasc is intermittent in its orrurrence. It is characterised
by a sudden wilting of the entire foliage, an oval hollow cavity in the crown and a
ba<ily difM*ased tap r(N>t which hitvi^s as a channel of inf«*ction.
Bacterial heart wilt. The bacteria attack only the innermost, tender leaves caus-
ing a wet, carlKtnaceoiiH rot. The organism has l>een isolated and its pathogenicity
proved.
1917] New Yobk Mseting 65
A bacterial blight of say bean. A. G. Johnson and Florence M. Coebpbb
For a number of years this disease has been under investigation at Madison, Wis-
consin. A malady apparently the same has also been reported from other parts
of the United States. At Madison the disease has been common during the past
three years, especially on the leaves.
These leaf lesions are small, rather angular spots, in late stages, dark in color,
brown to purplish black. In the earlier stages they are translucent and water soaked
in appearance and yellowish to light brown in color. The lesions may be irregularly
scattered or variously grouped and they not uncommonly coalesce. Rather incon-
spicuous glistening films of exudate are frequently noticeable on the lowfr surfaces
of the lesions.
Repeated isolation cultures have yielded a characteristic, white bacterial or-
ganism which has proved pathogenic on soy bean, producing characteristic lesions
as described above. The same organism has been reisolated from such lesions and
its pathogenicity in turn proved. This organism is a rod with rounded ends, motile
by a single polar flagellum, hence referable to the genus Pseudomonas of Migula
or the genus Bacterium of Ehrenberg as interpreted by Erwin F. Smith.
Studies on the physiological characteristics of the organism and its pathogenicity
on other leguminous hosts are in progress.
Host limitations of Septoria Lycopersici. J. B. S. Norton
Inoculations of seedlings of a number of Solanaceae and eighty varieties of tomato
in humid enclosures, with Septoria from tomato resulted in infections on several
species of Solanum, eggplant, Datura tatttkif and especially on potato, currant tomato
and Solanum carolinense. Spots developed better and spores larger on potato and
horse-nettle than on tomato, while the Datura spots were slow-growing, light colored
and small-spored. With larger plants outdoors, infections appeared rarely except
on Lycopersicum ; but undoubted infections resulted on horse-nettle and potato
and occasional pycnidia developed with spores smaller than on tomato. The to-
mato varieties in the seedling stage, showed decided differences in susceptibility
to the Septoria, both in number of infections and time of development. Many
dwarf varieties showed marked susceptibility.
Wintering of Septoria petroselina var. Apii. W. S. Krout
Heretofore, the seed has been considered the primary source for dissemination
and wintering of this organism. The following observations and results of tests
indicate this is not the case: (a) Pycnidia with spores are found on the pedicles
and have been reported on seed, (b) All spores taken from the dried pedicles
failed to germinate, (c) Spores from green celery tissues subjected to dessication
for eight months under laboratory conditions failed to germinate, (d) Young
seedlings in the seed-bed were never infected unless planted on soil that had pre-
viously grown celery infected with this organism, (e) This organism forms sclero-
tial-like intercellular bodies in the petioles, (f) Celery grown in new localities
gradually becomes infected, (g) Seed from the same bag, but divided and sown
upon two different farms produced the diseased seedlings in one case and healthy
seedlings in the other.
These studies indicate that the disease is not carried in the seed but in manures
containing diseased, decomposed plants, and probably by other methods.
Laboratory work has shown that heating celery seed to 50**C. for one-half hour
will eliminate all chances (if there be any) of the disease being disseminated through
the seed and pedicles.
Incomplete studies on Cercospora Apii Fr., indicate similar conditions.
66 Phytopathology [Vol. 7
A nematode disease of the dasheen and iU control by hot water treatment. L. P. Btaks
During the summer of 1914, a new disease of an economic aroid, the dasheen,
Coiocasia esculenta (L.) 8chott, waa found at one point in Florida causing serioua
damage. The malady is caused by the widely distributed nematode or eel worm,
Hcterodera radicicola (Grcef) Moller, which causes root-knot of many wild and cul-
tivated plantK, but which has not heretofore been authentically reported on the
dashren. In some places it has caused almost a complete failure of the dasheen
crop and is roffardcd as the most serious pathological factor in the successful pro-
duction of thiH plant.
The dit4asp has been found on dasheens in most of the South Atlantic States where
€M*onoiiiir aroids arc grown and in a shipment of cormcls imported from Egypt for
propagating pur]>08os.
On <lashcon roots the nematode produces macroscopically conspicuous svi'cllings
which hinder normal absorjition. On the surface of the tuberous growth it causes
protuberances and definite raiscMi areas resembling open sores, through which s€M?ond-
ary hel<l and storage parasites may readily enter. The eelworm docs not live on
the aerial parts of the <lashet»n. but, in rase of severe infection, it causes these |>or-
tions to be greatly r<HluriMl in size, and gives to the plant as a whole a decidedly sickly
app«'aranec\
The disease has been surressfully controllcHl by planting on uninfecte<l land se-
lected rormels from disrase-free areas, or diseaswl connels which have been treatetl
with water at o()°(\ for forty minutes.
Sot* worthy }*orto Rirau plant di^icaxcn. V. L. ST*:vE.\rt
To U' printed in full in thi* April issue of Phytofathouxjy.
Suljuriruj Concord grnjHH to prevent jHncdery mildew. F. K. CiLADWIN and Donald
Hkoouk
Continuing work rcfxirtcHl in Internat. Cong. Vit. Off. Kept. 1916: 117-12o. 1916,
plats of Concord grape vines were duste<i three times, July 18, August 2 and August
16, with sulfur-Iinie mixtures containing respectively twenty-five, fifty and seventy-
five percent »iulfur flour, ninety-five per cent or more of which pa-sses 'J(X)-mesh sieve.
The mixtures were applieil at the rate of forty pounds per acre. A single applica-
tion of liordeaux mixture was made <»n one plat on August 11. Treatinl plats al-
trrnuted uith rherk plats and all plats were separatnl by one buffer row.
I*o\i<lerv mildew, causetl bv I'nrinula neeator, was abundant. At harvest time
f»ne untreat<Hl plat showetl four per rent of the rlusters free from mildew and an-
other only tl.U()7 per cent fr<*e. The lM)rdraux-!<pray<'<i vines showi*<l six per cent of
th<* elu>«ti*rs free from mildew, the mixture rontaining sfventy-five per cent sulfur
showeil ninety-six j)er«M*nt frer. that containing fifty per cent showtn! eighty-three
per cent fr<*<* and that containing twenty-five per cent sIiow^hI twenty-seven i>er
cent free. The seventy-five per cent mixture causi»<l severe burning, the fifty per
cent mixture a small amount of biiniing and the twenty-five per cent mixture slight
burning
The nrrial triage of (\dtonfMtrium elephiintoiHtdin. (»Eo. (I. IlKUCK^tK'K and W. H. Ix)SO
Voung triM»s of l*\nuA heterophylUi in the greenhouse at Washington. D. C, were
in<HnilAti*d under cc»ntrolli*<i conditions in November. 1915. with the teliospores of
CtUrtn^fHtnum flrphnntofMHiiM (Schw./Thnm. In February, 1910. the aecia of Pcri-
dermtum carneum {lUtBc.) Seym, and Karle appeareti on the needles in abundance.
1917] New York Meeting* 67
These were fully mature late in March. Inoculations with the aeciospores March
7 and April 5, 1916, on the leaves of plants of Elephantopus tomentosus L. produced
in abundance the characteristic uredinia and telia of Coleosporium elephantopodia.
During the past three years parallel sets of inoculations of plants of Vernonia
on the one hand and of Elephantopus on the other with the aecia of Pertdermium
carneum from a number of species of pine have resulted in producing Colesoporium
vernonicB B. and C. on the former, and C. elephantopodia on the latter, indicating the
identity of the two species.
Pertdermium carneum is now reported for the first time on the needles of Pinus
carihcBa Morel., P. clausa (Engelm.) Sarg., P. echinata Mill., P. glabra Walt., P.
heterophylla (Ell.) Sudw., P. ponderosa Laws., P. rigida Mill., P. scopulorum (En-
gelm.) Lemm., and P. serotina Michx.
The Peridermium belonging to Coleosporium ipomceoB, George G. Hedgcock and
N. Rex Hunt
Peridermium ipomceoB a new foliicolus species on Pinus echinata Mill., P. palus-
tris Mill., P. rigida Mill., and P. taeda L. is described, with a range from Pensylvania
to Florida and Texas.
Plants of Ipomcea lacunosa L., /. pandurata L., /. triloba L., Pharbitis barbigera
(Sims.) G. Don., P. hederacea (L.) Choisy, and Quamoclit coccinea (L.) Moench un-
der controlled conditions were successfully inoculated with the aeciospores of this
Peridermium, producing on their foliage the typical uredinia and telia of ColeospO"
rium ipom(EXB (Schw.) Burrill, thus proving that it is the aecial stage of this Coleo-
sporium. Plants of species of Amsonia, Aster, Calonyction, Chrysopsis, Convol-
vulus, Coreopsis, Elephantopus, Helianthus, Laciniaria, Silphium, Solidago, Ver-
besina and Vernonia were unsuccessfully inoculated.
Coleosporium ipomcece is now reported for the first time on the leaves of Ipomolea
caroliniana Pursh., 7. trifida (H. B. K.) G. Don., and Pharbitis barbigera,
A Peridermium belonging to Coleosporium terebinthinacece. Geo. G. Hedgcock
and N. Rex Hunt
Peridermium terebinthinaceumf a new foliicolus species on Pinus echinata Mill.,
P. rigida Mill., and P. taeda L., is described with a range from North Carolina to
Georgia.
Inoculations were made under controlled conditions with the aeciospores of
this Peridermium on plants of Silphium asteriscus L., S. intcgrifolium Michx. ^ S. tri'
foliatum L. and Parthenium integrifolium L. in May and June 1916. In about two
weeks the uredinia, and later the telia of Colesporium terebinthinacea; (Schw.)
Arthur appeared on the leaves of all these species, proving the Peridermium to
be the aecial stage of this Coleosporium. Inoculations were made at the same
time on plants of species of Amsonia, Coreopsis and Laciniaria without result.
Coleosporium terebinthinacece is now reported for the first time on the leaves of
Silphium angustum (A. Gray) Small, S. compositum Michx., S. dentatum Ell., S.
glabrum Eggert, and S. pinnatifidum Ell.
An alternate form for Coleosporium helianthi. George G. Hedgcock and N. Rex
Hunt
A new foliicolus species, Peridermium helianthi, is described on Pinus virginiana
Mill., with a range from Pennsylvania to South Carolina and Tennessee. Inocula-
tions made with the aeciospores of the Peridermium, under controlled conditions,
Phytopathology [Vol. 7
on planU of HelianthuM decapetaluM L,, H, divawieaiui h,, H. ffiganteuB L., H, gla^mu
Small, and H, hir9utu9 Raf . produced the uredinia and telia of CoUo^porium kelianiki
(Schw.) Arthur, usually in abundance, thus proving the Peridermium to be the
aecial at age of this Ck>leo8poriunk. Inoculations were made at the same time on
plants of species of Aster, Chrysopsis, Coreopsis, Elephantopus, Laciniaria, Parthe-
nium, Rudbeckia, Silphium, Solidago, Verbesina, and Vcmonia with negative re-
sults. The results of these inoculations indicate that the Coleosporiums on Coreop-
sis and Verbesina arc distinct from the one on Helianthus. The Coleoeporium in
Florida on Verbesina which has been assigned to C. heliarUki may belong to one of
the unnamed species of Peridermium found by the writers in the south.
CoUo9ponum heliantki is now reported for the first time on HelianthuM au9traii9
Small, //. divaricatus L., //. eggeriii Small, //. glaucus Small, H. gro99€'9erraiu9
Martens, //. hirsutus Raf., //. microcephalus T. and G. ,and //. aaxicola Small, and
its range (on Helianthus) is extended to Louisiana and Florida.
SoffM fi^tr ha$U for CoUo9parium »olidagin%$. Georqe G. Hedocock and N. Rxx
Htnt
Peridermium acirolum I'ndcrw. and Earlc, the aecial stage of CoUoBpcrium coli-
daginia (Schw.) Tht'im.. is reported for the first time on Pinun carihaea Morel., P.
coniarta Loud.. P. diraricata (Ait.) Du Mont do Cours, P. echinata Mill., P. tUioltii
Kngolm., P, mayriana Sudw., P. nigra Arnold (P. laricio Poir.), P. nigra var. ays-
iriaca Srhneid., P. jtaluatria Mill., P. ptpnderoaa Law8., P. acopiUarum (Engelm.)
liPmm., P. nrrotina Mirhx., P. taeda L., and P. thunhergii Pari.
Positive results have been obtained from inoculationfl with the aeriospores of
Peridermium acirolum on plants of species of Aster and Solidago, and negative re-
sults only, on plant** of Hpc*cics of Campanula, ConvolvuluK, Coreopsis, Elephanto-
pus, Kupat<irium. Euthaniia, Helianthus, Ip<>nH>ea, Laciniaria, Parthcnium, Phar-
bitiK. iS'iH'i'id. Verbesina. and Vcmonia. A Coleosporium on Chrytopaia mariana
(L.) Ntitt.. found firMt by \V. H. Ixing in Florida, in tentatively assigned to this
sperics.
The range* of the Peridermium has be<»n cxtendwi to include nearly all States from
Minnesota and New Hampshire on the north to Arkansa.H and Florida on the south.
Sotrf on name tijterirn of ColroafMtrium. (iKr>K(2K (i. HKDttriK'K and N. Rkx Hunt
ColronjHtrium delicatulum .\rth. and Kern is report (h1 for the first time on Euthamia
earoliniaua (L.) (ireene, and E. leptorephala (T. and (•.) Greene, and Peridermium
delicatulum (.-Xrth. and Kern) Ilfnlge. and l^uig for the fir<«t time on Pinua tariteif
Morel.. /•. echinata Mill.. P. eUiottii Engelm.. P. hrUrophylla (Ell.) Sudw., P. may-
riana Sudw.. P. palimtrin Mill.. P. nigra .\rnoM. /*. /tonderosa Lawn.. P. reainoaa
Ait.. P. Htrotina Mirhx., ami /'. tarda L.. and the range of the spei*ii*s is greatly
e\tend«l.
CotfoffHtrium laciniarnr .Vrth. i.M reiM)rt<*<l for the first time on l^aciniaria earlei
(•reene. L. drgann (Walt. ) Kuntze. L. ttongata (tre<»ne, L. /xtuciflora (Pursh) Kuntie,
L. acariima < L. i Hill, and L. utrotina (!re<>ne, and the range of the specie's extended
to Klnridu on the .^fiuth and to New Jersey un the north.
Some ri« 1/ hit<*t)* ftn ('itl*tt*fmn turn inritntpicuum. (iKoHciK (i. Hk<iD<'<x*K and N. Rex
Hi .ST
Ac*<-io*>|Niri*s fr«»ni PiTxdtrmium tnconapicuum I^mg rollert4*d for the first time on
the nc4*4lles of Ptnuf echtnata Mill, were Huce<*ftsfully inorulat<*d on the leaves of
1917] New York Meeting 69
both Coreopsis major omUeri (Ell.) Britton and C. verticiUcUa L. resulting in the
fonnation of the uredinia and tellia of Coleosporium inconspicuum (Long) Hedge,
and Long. Unsuccessful inoculations were made on plants of species of Amsonia,
Aster, Chrysopsis, Elephantopus, Euthamia, Helianthus, Laciniaria, Silphium,
Solidago, Verbesina> and Vemonia. The results of these inoculations indicate
that Coleosporium inconspicuum is distinct from species found on plants of these
genera. This Coleosporium is reported for the first time on Coreopsis delphinifolia
Lam., C lanceolata L., C. major Walt., C. major rigida (Nutt.) Boynton, and C.
Iripteris L.
Coleosporium apocynaceum Cooke has been collected at Clearwater, South Caro-
lina on Amsonia cliate Walt.
•
Scolecotrichum graminis on timothy, orchard grass, and other grasses. A. G. John-
son and C. W. Hunoerford
During the past few years Scolecotrichum graminis has been observed by the
writers on timothy and orchard-grass at various points from Wisconsin to the Pacific
Coast. The fungus causes a serious disease of these hosts especially in Wisconsin.
The young lesions on the leaf are circular to elliptical in form, vary greatly in size,
and are usually purplish brown in color, The older lesions turn grayish brown as
the invaded tissues die. In severe cases they coalesce involving considerable por-
tions of the leaf-blades. In the worst cases practically all of the leaves of affected
plants are dried up at about flowering time. In moist weather the fungus sporulates
abundantly on the older lesions. On orchard grass, the lesions are somewhat more
distinctive and sporulation takes place more abundantly. The conidiophores come
out through the stomata and form dark-colored tufts arranged rather regularly
in rows.
Other grasses observed by the writers as hosts for the fungus are Agrostis alba,
Bromus margimitus, Bromus sitchensis, Hordeum jubatum, Hordeum nodosum, Ely"
mus glaucus and Elymiui robustus.
Observations at Madison have shown that the fungus over-winters readily in
tufts of orchard grass and timothy and resumes activity early in the spring.
Bacteria of barley blight seed-borne, L. R. Jones, A. G. Johnson, and C. S. Reddt
* In further studies on the bacterial blight of barley, upon which reports have
been previously made, the mode of overwintering of the causal organism and of its
dissemination over long distances have received especial attention. Field evidence
early indicated that in certain cases the disease doubtless was introduced with
seed from various western sources. In following up this matter, seed was collected
in 1914 from a field of barley in Montana severely affected by the blight. Lesions
were evident on the glumes of these plants before maturity and showed, although
less clearly, upon the ripe grain. Some of this seed was planted in our trial grounds
in Wisconsin in 1915, and the blight developed abundantly upon the leaves of the
young plants.
Isolation cultures were made in July, 1916, from the glumes of barley kernels
from this same 1914 Montana collection. The characteristic barley blight organism
was obtained and its pathogenicity proved by inoculation experiments. It is thus
apparent that the organism may be carried with the seed grain and remains viable
after at least two years of dormancy. Preliminary trials indicate that the organism
may be destroyed by seed disinfection.
70 Phttopathologt [Vol. 7
The Ptteudopeziza leaf spot diseases of alfalfa and red clover, Fred Reucl Jonks
The PseudopeEiEa leaf ftpots of alfalfa and red clover have been studied for the
past two yearn for the purpose of determining the following points.
1. Are the causal organisms the same or distinct species?
2. Is any other spore-form than the ascospore included in the life history of these
fungi?
3. What is the relation of these fungi to the tissues of their hosts?
4. How do these fungi oven»inter?
5. Can the occurrence of the alfalfa leaf spot on alfalfa sown in a new region for
the first time he prevented by seed treatment?
Progn^s hat* been made as follows:
1. Both fungi have been obtained in pure cylturc. Slight morphological and
distint't phyniologiral differences have been found.
2. Only ftf^cosporcs have been found produced in nature. Conidia-like structures
occur in cultures.
3. (icnninnting aMcospores penetrate the epidermal cells directly and the mycelium
devt'lopM within the ho»t ccIIh and penetrates the cell wuIIh.
4. The fungtiH <)ver>»inters on dead leaves which escape <lecay, and ascospores
developiMl either in old or new apothecia are a source of spring infection.
5. .Mfalfa serd very th(»roughly (iisinfcrt«*<l has been sown at distances up to 15
miles from other alfalfa. Leaf spot has occurred on all these plots.
The development of the aeciid Mtagc of \igredo on red rlorer. I. K. Meliits and Wil-
liam DiKHL
Th«* orrurrence of the ure<liiio- and teli<»spore stages of Sigredo fallens (Desm.)
Arthur \l'rom\fetn fallem* (I)esni.) Kern) on red clover {Trifolium pratense) is com-
mon, but the aecial stagt* of this nist is apparently not well known. The ure<iino-
spore stage di'velopi**! abundantly on re<l rlover growing in the greenhouse l>egin-
niiig Hb<»ut Januarx 10. VJU), when ihe clovei plantswere alniut six inches tall. These
grew in pots forming a border nearest tin* glass on a bench in a house where the
temprrnturr rang(*il from 10" at night to '2(f(\ in the day time. On .March 26,
Il»l*>. a«MMa Urn* nlisrrv<*<l <»n the leaves of the red cIovit plants. During the next
two uf«*ks the aerial stag** breaine abundant. It rontinued to develop for about
a month whrti the tmnpcrature raised in th<* h(»use due to the increaseti sunshintv
Whit*' rIovMT \Trift'hum refnnf*), alsike ( 7\ hifhridumi, and crimhon cl<»ver (7*. i«-
earnatumi. growing ui close proximity were c<intinually free fnmi infection.
H<-p«*at(il att<'mpts were niadt* to transfer the rust to the above hosts by using
the acrio?.pon'f« but no infections were obtained except on the re<i clover. It would
appear that .Vi*;r«#/n/*i//« n/* is autm*cious and not lieter(K'cious as h<*retofore reportwl.
.4 rnoluutrittnu timtaite of the Irish jnttnto and itf< rnntrtd. II. .\. Kl)HO.\ and Oswald
SlIHIllNKU
liegitiiiing eariv in the past July, potato plants in numerous eastern fields from
Mauie to N'irginia developed a downward curling of the leaf margins accompanicNi
by a bronzing and later a brriwning but not a yellowing of the foliage. Death of
the leaves and sudden collapse of the stems at the ground level follow<Hl. Fungi
of parasitic habits a|>peared at and above the |M>int of collapse. Dr. W. A. Orton
c»b»erved the disease in New Jersey an<l note«l its absence fn»m areas treated with
fM»tn«>h or stable manure. Similar obsi'rvations were made elsewhere. partieuUtrly
in Mame. where Dr. Joseph l{os4*nbaum and B. K. Hrown ami L. A. Hurst have
1917] New York Meeting 71
undertaken a study of the disease in cooperation with the Maine Agricultural Ex-
periment Station. These investigations are incomplete but present indications
are that the fungi isolated are weakling parasites operating as contributing factors
and that the primary cause is malnutrition, resulting from insufficient potash or
perhaps an excess of nitrates in the presence of a minimimi potash supply. In Maine
the disease appears to be correlated with certain soil types and is most marked on
Irish cobbler, Bliss and Eureka, though not entirely restricted to these early varie-
ties. Stable manure is an excellent corrective.
Notes on curly dwarf symptoms on Irish potatoes. W. L. Durrell
Plants showing curly dwarf symptoms were very prevalent in Iowa this past
season on the varieties Irish cobbler. Rural new yorker and Early ohio. In some
cases these symptoms were on plants grown from the progeny of plants having shown
curly dwarf symptoms the preceding year; in others they were induced by climatic
conditions. The disease made its appearance on the early planting about June 10
and continued to develop throughout the season. In August, plants that had been
normal up imtil that time, showed typical signs of curly dwarf on the foliage pro-
duced during this period. The upper third of the plant had shortened internodes,
crinkled and curled leaves, giving this portion of the plant a bushy appearance so
characteristic of curly dwarf. These symptoms were induced in the field by the
hot dry weather during August and similar ones were later artificially developed
in the laboratory. The plants manifesting these induced symptoms of curly dwarf
put forth normal foliage again in September with the advent of cooler weather.
Histological studies show that the crinkling of the leaves is due to necrosis of
certain epidermal and cortical cells of the veins, followed by the growth of the par-
enchyma cells which induces a buckling of the leaf surface. In addition the leaves
showing curly dwarf symptoms were characterized by a most marked decrease from
the normal in the size of the parenchyma cells. Furthermore transpiration tests
using the cobalt chloride paper and potometer methods indicate that curly dwarf
plants transpire more rapidly than normal ones.
Notes on mosaic symptoms of irish potatoes. I. E. Melhus
The so-called mosaic disease of potatoes is characterized by yellow mottling and
crinkling of the foliage. Its effect on the potato plant, transmissibility, and rela-
tion to curly dwarf is very imperfectly understood. The varieties of Bliss triumph
and Green mountain, which showed mosaic symptoms in 1914 and 1915 in northern
Maine, were planted in 1916 at Ames, Iowa. The characteristic yellow mottling
typical of this disease did not develop at any time during the growing season. Curly
dwarf symptoms, however, were prevailingly present.
(a) Plants seemingly badly affected in 1915 produced 0.237 pounds per hill.
(b) Plants seemingly moderately affected in 1915 produced 0.29 pounds per hill.
(c) Plants seemingly slightly affected in 1915 produced 0.32 pounds per hill.
Healthy plants used as checks produced 0.46 pounds per hill.
None of the plants in lots a or 6 grew as large as those in lot c, but some of the
c-plants were like those of a. Although the typical mottling characteristic of mo-
saic in certain northern potato growing districts may not develop, the progeny
of plants showing these symptoms are undesirable for seed purposes.
Frost necrosis of potato tubers. L. R. Jones and Ernest Bailet
A peculiar type of non-inheritable "net necrosis" of potato tubers has been under
observation for several years under conditions suggesting frost injury. Carefully
72 Phttopatholoqt [Vol. 7
repeated ohilliog experiments confirm this hypothesis. Tubers ''frosen toUd" are
totally killed and collapse when thawed. If, however, the chilling stops with in-
cipient ice-crystallisation, killing may be confined to such interior tissues as are
most sensitiye. Such chilled tubers appear normal externally but when cut show
the interior vascular regions to be most sensitive and hence the first to succumb
and discolor. Therefore, moderate exposure to freesing temperature may pro-
duce either ''ring" or "net" necrosis, the blackened vascular portions permeating
the starchy fundamental tissues. Individual variations in sensitiveness occur be-
tween tubers, but in general the best t3rpe8 of ''net necrosis" have been secured
by about two hours exposure to +5*C., with similar results by exposures ranging
from — l^C. for 8.5 hours to — 9*^0. for one hour. Slightly more severe treatments,
or unequal exposures may give frosen spots with corresponding dark blotches in-
volving the general parenchyma. The stem end of the tuber is always more sensi-
tive than the other.
Will Spongoapora subterranea prove serious in Virginia? J. A. McCuntock
Potatoes affected with powdery scab planted in Virginia in the spring of 1916
produced a crop free from this disease. In the spring of 1916 affected tubers from
Maine were planted at Norfolk, Virginia, and at Tasley, on the Eastern Shore of
Virginia. The writer examined the crop at each place but found no signs of Spon-
gospora infection on any of the harvested tubers. Some of the infected seed from
Maine was held over summer in cold storage and planted at Norfolk about the
time the fall crop of Irish potatoes was planted in Virginia. On November 23, 1910,
the tubers were harvested and examined but no Spongospora infection was observed.
These results corroborate those of 1915 and lead one to conclude that powdery
scab will not be prevalent either on the spring or fall planted potatoes even though
the seed tubers are infected with Spongospora subterranea.
Seed potato certification in Nova Scotia. Paul A. Murpht
Many fields of Garnet chili potatoes for the Bermuda seed trade were infected
to the extent of fifty per cent with leaf roll, and whole districts to not less than ten
per cent, making a difficult situation when we took charge. Hill selection in dis-
tricts proved useless in several cases tried. The scheme adopted had to be a com-
pn»hensive and unusual one, an outline of which follows.
It is neccHsary for growers to start with stock of good previous record. This
provision, which is absolutely insisted on, is becoming more stringent, and in two
years the growers will use exclusively stock which is now selected and is being given
a three years' trial. Thereafter it will still be continuously selected, one man be-
ing ap|H)intecl in each district to grow it for his ncigbors.
The munriier inspection iM made jointly by officerH of the Departments of Agri-
culture of (*anada and Bermuda, while the autumn inHpection is made by the Cana-
dian aiithoritii*?*. The grower'n name appearH on each barrel, whereby many stocks
are trurcd to the Bennuda pint at ions. Am a further nafeguard a sample of every
gruwcr'M potatoes are ^eiit to Bennuda, where they are all plante<l together.
The fcornfmw importance of mosaic of potato. Vm'L \. Mukphy
The b<*ginning of a serii»« of experiments to inve«tigate the economic importance
of iUin dineaite han given striking resultn. In one ex|K*rinient of eleven similar plots
Hide by Hide, plaiit^Ml partly with ht^althy, and partly with mosaic-diseased Green
mountains of the name strain, the dim^aiuti plants gave a yield which was on the
1917] New York MEBTma 73
average only 57.8 per cent that of the normal plants, the limits being 52.0 per cent
and 63.6 per cent. Furthermore the crop of the mosaic-diseased plants was mar-
ketable only to the extent of 82.7 per cent (limits, 74.1 and 87.3), while 91.6 per cent
of the crop of the healthy plants was marketable (limits, 81.9 and 94.1). This means
that in an average crop of 300 bushels there is a loss of one and one-third bushels
of marketable potatoes for every 1 per cent of mosaic present. This coupled with
the fact that the trouble is constant every year and that it generally affects, where
present, not far from twenty per cent of the crop means a steady loss of from twenty
to thirty bushels of potatoes per acre per year. The eating qualities of the pota-
toes are not impaired.
A new strain of Puccinia graminis, E. C. Stakman and F. J. Piemeisel
A rust which behaves differently from any of the common biologic forms of Puc'
cinia graminta has recently been found on club wheat and a number of wild grasses.
It resembles P. graminia tritici morphologically and parasitically more that it does
any other biologits form. However, the common Trittcum vtUgare wheats which
have been inoculated are highly resistant to it. Both P. graminis tritici and the
new strain have a niunber of hosts in common, viz: Triticum compactumf Barley,
Agropyron smithiif Elymus canadensis, Elymus macoum't, and Hordeum jubatum.
The new strain has also been found in nature on Elymus glattcus and E, condenscdus
and has infected a number of grasses in the greenhouse. Extensive cross-inocula-
tion experiments are now under way.
The rust was found only west of the Rocky Mountains in Idaho and Washing-
ton where it seemed to take the place of ordinary P. graminis tritici, none 6i which
was foimd in the region mentioned.
Puccinia graminis on wheat kernels and its relation to subsequent infection. Chab.
W. HUNGERPORD
Various workers have noted the occurrence of rust pustules on seeds of different
grains and some have held that the fungus might infect the plant by this means.
Experiments have been carried on at Madison, Wisconsin the last year to deter-
mine if possible whether Puccinia graminis is able to infect wheat through the seed.
Three lines of attack have been followed, (a) Rusted seed after being germinated
at different temperatures has been fixed and examined by histological methods and
in no case was the fungus found to penetrate the embryonic tissues, (b) Treated
and untreated samples of rusted seed, as well as clean seed, were planted in the
field and the first appearance of stem rust upon the plants in the different plots
was noted at practically the same time, (c) Two lots of rust-infected seed have
been grown to maturity in an isolated room in the greenhouse. No rust has ap-
peared on any of these plants. Although the work has not been fully completed,
the results so far tend to show that seed wheat infected with Puccinia graminis
does not cause infection of the wheat plant.
Similar experiments are being started at Corvallis, Oregon, with wheat infected
wi th Ptu:cinia glumarum.
Ecological observations on Ustilago Zeae. Alden A. Potter and Leo E. Melchers
Pammel and Stewart in 1893 observed that the nodal buds of maize were particu-
larly subject to smut and that "where one smut boil made its appearance on the
lower nodes, others appeared further up.'' It thus becomes desirable to explain
how the infection, shown by Brefeld to be strictly local in its development, can
spread on the plant. The basis of study has been Brefeld' s idea of distribution by
74 Phytopathology [Vol. 7
air-l>orne conidia. The organism han boon iflolated in pure, conidial culture, both
from the air and from the young plants some little time before the disease appeared.
The com plant is thus seen to be well adapted as a spore trap. The conidia caught
probably do not infect directly. The result is rather the development of a virulent
culture in the leaf axil. A plant may thus become a center of aerial distribution;
or, when rain recurs, the conidia may be washed down or splashed out upon other
leaves. Thus it may sometimes happen that all the culms of a plant, or hill, will
show many no<lal infections when an equal number of stalks immediately adjacent
will not be infecte<l at all.
The shart-cyrUd Vromycen of Xorth America. G. li. Hishy
Only eleven species of short -<*yrle<l rnmiyces huvc boon found in Xorth America.
Those nists are parasitic upon six families of Monocotylodons and Dicotyledons.
Various relationships are evident botwoon those rusts and other long-cycled and
short-cycled species of ruMts. Seven sperios arc commonly micro-forms; for four
of th(»so, pyrnia are known. Seven sperios have strictly local mycelium. These
nists occur mainly in WcHtcrn and Southern North .Vniorira. The specimens have
been studied at the Arthur HerbarUim.
Holed and mycorhiza uf)on forest trees and an unusual mycorhiza ujwn trhite oak.
L. H. Pe.vxington
One instance of a Boletus, li. nperiosus Frost, connect o<l with mycorhiia of oak
was reported in VJilH. Since that time five other species, li. froHtii Kussell, B. in-
deMiitu* I*k. li. chroma ften Frost, li. purpureus Vr. and H. gracilis Pk. have been found
conn«»ct<»<l with mycorhiza of forest trees, usually oaks. Two of these species,
a. froftiii and H. indecisun, have been found to produce sclerotia similar to those
report e<l f<»r li. sfxciimuM.
A peculiar form of mycorhiza was found upon white-oak roots in which the hy-
p<Ttrophie<l branches are closely aggrogateii and surrounded by a peridium-like
layer of fungal tissue. This gives them the appearance of white root tubercles,
four to twolve millimeters in <liameter. These tuborcle-liko growths are not unlike
thos<» uiH)n bee<'h roolH d«»scribe<i in 1890 by Von Schronk. They are also definit<*ly
connoctecl with small sclerotia from which there in a growth of mycelium in early
summer to prcHluoe now mycorhiza U(Mm the roots. .\t tempts to inoculate the
nK)tH of oth(*r treses with this fungus have thus far failo<l.
A nrw paranitic nlime mold nuitahle for class tatrk. John A. Klliott
The Hwoot [lotato "|m>x" organi-nm, Cystosfntra haiata Klliott, as it occurs on sweet
|x>tftt<M'?«, offors itself an an oxrollont «*xainplo of the Plu.*<mo<liophoral(*s for use in
tho Inbonitory. Infort<'<l i>IniitM growing botwoon shoots of moist blotting paper
afTnrd »ii abundatiro nf pnriu«itizo4l rootlets and growing i>oints of stems for free-
hand !*«'<>! ioiiiiig or for oiiibo<idiiig in paraffin. Such matorial iH easily sectioned
and rontaiii^ groat numborM of tin* para>ito in all stag<'H of its life cycle. The rapid-
ity uitli \iliirh tho organism goos through its complete life history makes a study
of living nmtoriul of ^pecial value.
Straiuf nf Hhiztuiimia. J. RosKMiAt'M and M. Siiapavalov
During tho mimmor of llll(> a strain of Hhizootonia was isolated from potato steins
which showinl a girdling and hollowing at or near the surface of the ground. This
1917] New York Meeting 75
strain, designated R 5, differs in the following particulars from other strains isolated
from stems and tubers of potatoes grown in Maine and Florida:
(1) Inoculations with R 5 produced definite lesions in injured potato stems grow-
ing in the field and greenhouse and injured tubers, while in the case of inoculations
with other strains the lesions, if produced at all, were smaller and the results not so
conclusive. The injured checks remained healthy.
(2) Macroscopically R 5 can be distinguished by the darker coloration of the
medium, especially when grown on potato agar, and by the light grayish sclerotia
as compared with the dark-brown sclerotia of the others when grown on corn-meal
agar.
(3) Microscopically R 5 differs from the other strains in its finer mycelium, w>>ich
measures 5 to 9 m in diameter while the others measure 10 to 14 /x.
Is it not possible that different strains of Rhizoctonia may offer an explanation
for the conflicting reports regarding artificial infection?
The aerial stage of the red clover rust. W. H. Davis and A. G. Johnson
The well known red clover rust, Uromyces fallens (Desm.) Kern, has long been
suspected of having an aecial stage. Our observations and experiments have thrown
definite light on the question.
Aecia on red clover were first obtained experimentally in the greenhouse in Decem-
ber 1915 and similarly again in January and February, 1916. Later in the spring
a number of cases of aecia were observed on the same host out of doors near Madison.
Following various sowings of aeciospores from such sources on rust-free red clover
plants under glass, uredinospores developed uniformly. These were identical with
those conmionly observed on that host. Sowings of viable teliospores resulted in
the development of aecia, identical with those observed in greenhouse and field.
It is thus evident that this rust is a long-cycled autoecious species, i.e., with
pycnia, aecia, uredinia and telia on the same host.
Observations on pear blight in Illinois. F. L. Stevens, W. A. Ruth, G. L. Peltier,
and J. R. Malloch
Experiments made by applying Bacillus amylovorous in suspension in water to
pear buds in 1915 did not indicate in 1916 that the bacilli hibernated in the buds.
Subcuticular infections of spurs from hold-over trunk cankers occurred in 1916,
with a maximum number of twelve such infections from one canker. The organ-
ism appeared to be dead in all twig cankers. A few living cankers provided exudate
for serious well-distributed blossom infection, which in turn provided exudate for
further infections, these continuing until early in June.
Leaves appeared to be at no time naturally infected from the exterior and on
June 1 blades and pedicles could not be inoculated though the fruit and pedicles
were still susceptible. Bordeaux mixture controlled the floral infection without
reducing the set of fruit.
Second progress report on investigations of leaf spot of cherries and plums in Wisconsin.
G. W. Keitt
Comparative studies of Coccomyces hiemalis Higgins and related organisms in
connection with leaf spot diseases of cherries and plums have been continued, and
spraying and sanitation experiments in the control of cherry leaf spot have been
begun. Only the control work is reported here.
Spraying (Montmorency and Early Richmond). In early summer, the disease
occurred in unusual severity, but, after the advent of hot dry weather in late July,
76 Phttopatholoot [Vou 7
it made relatively little progress. It was satisfactorily controlled by Bordeaux
mixture, 4-4-50, 3-3-50, and 2-2-50, and lime-sulphur (commercial ooncentrmte,
33*B.), 1-10, applied (1) when the petals fell, (2) l(m7 days later, and (3) just after
the fruit was harvested. "Atomic sulphur," 5-50; barium-sulphur, 3-50; and aelf-
boiled lime-sulphur, 8-8-50, in parallel applications, did not control the disease
satisfactorily. An additional application just before the blossoms opened did not
increase the efficiency of leaf spot control.
Sanitaiian. Life history studies, sanitation experiments, and extensive obeerva-
tions indicate that, under Wisconsin conditions, the spray schedule may be strongly
reinforced by turning under the fallen leaves as completely as feasible by clean cul-
tivation before the blossoms open (In 1016, the first ascospore discharge was observed
as cherry blossoms began to open).
Jonathan spot, Charles Brooks and J. S. Collet
The development of Jonathan spot increases with an increase in temperature up
to 20*C., but is entirely inhibited at dO'^C. The disease can be readily produced
in saturated air in closed moist chambers but seldom develops in a stirred air of
70 per cent or 95 per cent relative humidity.
Temperature relation* of apple rot fungi. Charles Brooks and J. S. Collet
Most apple-rot fungi will grow at a lower temperature on com meal agar than
on fruit and at a lowor temperature on ripe fruit than on green fruit. With several
if not all of the storage-rot fungi the initial stages of rotting are more inhibited
by low temperatures than is the germination of the spores. Rots may finally make
a fairly rapid development at temperatures at which the fungus is at first barely
able to make a start. Even at favorable temperatures most of the fungi pass through
a period of incubation on apples that is not evident on culture media.
Control of apple §cab by bleaching poicder. W. S. Brock and W. A. Ruth
Bleaching powder when applied to apple trees in 1016 reduced apple scab from 50
per cent to 11.2 per cent. No injury resulted to fruit or foliage. The leaves on
trees sprayed with this material were practically free from scab. Leaves on un-
■prayed trees showed serious seal) infection.
In 1010 the material was applied at high concentrations alone, and with lime,
causing little foliage injury, but no scab developed. Higher concentrations alone
and in combination with other materials will be tried.
Studie* on peach yello%c» and little peach. M. A. Blake, Mel. T. Cook and C. A.
ScHm-AKiR
S>'mptoms ver>' iiimilar to peach yellown and little peach may be due to other
causes. Tents with healthy and diHcaiMHl trees showetl (1) pulp from healthy leaves
retains original color longer than pulp from ditieased leaves; (2) juice from healthy
leaves in more mucilaginous than that from diseased leaves; (3) juice from fast-
growing trecM more niurilaginoun than juice from the leaves of slow-growing trees;
(4) leaves taken from healthv trees after sunset showe<l a minimum amount of
starch, leaves from little peach trees a larger amount, and leaves from yellows trees
and from girdle<l trees the greatest amount; (5) leaves from an apparently healthy
branch a<ljacent to a diseased branch on same tree showed a higher starch content
than leaves from a healthy tree; (7) leaven from fast-growing trees lose starch more
rapidly than leaves from slow-growing trees; (8) juice extracted from healthy
1917] New York Meeting 77
leaves showed less oxidase than juice from diseased leaves; (9) juice from healthy
kernels showed less catalase and acid than juice from diseased kernels; (10) the tan-
nin content of healthy fruit is less than that of diseased or forced fruit.
Pits from diseased trees failed to germinate. Budding experiments with dis-
eased buds indicate that the appearance of the disease in the young trees varies
with source of bud wood.
A Xylaria root-rot of the apple. F. D. Fromme and H. E. Thomas
A destructive root-rot of apple trees is prevalent in the chief orchard sections of
Virginia. The infectiousness of this condition is shown in the death of adjoining
trees in groups irrespective of soil conditions or topography, and in the death of
replants set in holes from which diseased trees were removed. Isolations from
affected roots from a number of orchards have yielded cultures of a fungus which
is apparently the conidial stage of a species of Xylaria. Perithecial stromata of
Xylaria polymorpha have been found on roots of apple trees showing typical attack.
Tjrpical root-rot lesions have been produced with pure cultures of the Xylaria in-
troduced into bark woimds of living apple roots in damp chambers and in the field.
The introduced fungus has been recovered in pure culture from these lesions. Two
or more species of Xylaria may be involved; their interrelations are subject fgr fur-
ther study. Apparently all varieties of apples are susceptible and probably equally
so. Observations indicate that the organism may be spread in cultivation, in the
removal of borers, in contact between root systems of adjoining trees and in sur-
face washing of infective material.
Pycnial acars, an important diagnostic character for the white pine blister rust. Regi-
nald H. COLLEY
After the pycnospores appear in their characteristic sweetish drops the whole
pycnium is cut out by a protective layer of tissue which forms at a depth of* several
cells below the pycnial layer. Everything above the protective layer drys out and
dies. The result of this drying is a rusty brown patch or scar which indicates by
its size the extent of the pycnium. The scars are dark brown and glossy at first.
Later they become rusty brown with a dry dusty-granular surface. They average
about four millimeters in diameter. Pycnial scars are a positive diagnostic field
character for the white pine blister rust, here reported for the first time. They
are especially valuable when the bark is but little swollen, and when there is no
indication of aecia or of aecial scars.
Mycelium of the white pine blister rust. Reginald H. Colley
The uninucleate mycelium of Cronartium ribicola forces its way between the bark
cells of white pine, frequently forming strands. As the cells are forced apart the.
bark s\^'ells. Haustoria penetrate practically every non-woody cell in the infected
tissue. The sieve tubes become plugged. Hyphae follow the ray cells past the
cambium and into the wood for a distance of at least three annual rings. Thus the
fungus derives nourishment from both the ascending and descending currents of
sap. The morphological characters of the mycelium are definite and constant un-
der all conditions observed. Therefore the mycelium has positive diagnostic value
for the blister rust before there is any exterior indication of spore formation.
The binucleate mycelium in Ribes leaves is limited in extent. Haustoria are rare.
Uredinia and telia form quickly and burst through the epidermis easily.
The binucleate mycelium is very abundant in cases of petiole infection and the
haustoria are larger and more numerous than in the lamina. Normal telia are
usually produced but sometimes they are formed internally.
78 Phytopathology (Vol. 7
A fpecien of Chrysomyxa new to North America, H. S. Jackson
The KpnuB Chrynomyxa was efltablished in 1840 by Unger with C. abietis (Wallr.)
Vng. aM the t>'pe spccieD, and has generally been interpreted as including both long
and short cycle forms. Arthur restricted this genus to include only the micro-
forms and established Melampsoropsis (Schrot.) Arth. for those species with a long
life cycle. All of the latter forms are assumed to be hcteroecious and have their
uredinia and telia on Pyrolaceae, Ericaceae and Vacciniaceac. The aecial stages
so far as determined, have proved to \tc species of Peridcrmium on Picea. In America
eight species have been reported, all of which are long cycle forms and are referred
to Melampsoropsis by Arthur. Four of these have been definitely connected through
cultures by European and American students with their aecial stages.
A short cycle form referable to the genus Chr>'8omyxa (as restricted by Arthur)
is recogniEcd in America for the first time and causes a disease of the leaves of Picta
engelmannii .
A Gnomonia on eggplant. C. W. Edgerton
During the past three years, a species of Gnomonia has been found on old egg-
plant stems during the winter season at Baton Houge, I^)uisiana. This fungus has
br<»n repeate<lly cultured and it has been found to be very similar, if not identical,
from a morphological standpoint, with the fungus causing the eggplant blight,
Phyltontirta hnrtorum. The oval Phyllostirta spores and the long narrow Phlyctaena
spores developc<l in culture. Cultures of the (inomonia and cultures of Phyllasticta
hortarum cannot be told apart. Inoculation experiments, however, have always
been negative. While it may be that the Onomonia has no connection with the
Phyllosticta, it is very probable that the two are closely related species.
LITERATURE ON AMERICAN PLANT DISEASES*
Compiled bt Eunice R. Oberly, Librarian, Bureau op Plant Industry
AND Florence P. Smith, Assistant
October to November, 1916
Avema-Sacca, Rosario. Molestias cryptogamicas da canna de assucar. Bol.
Agr. [Sao Paulo] 17, no. 8: 610-641, illus. Agosto, 1916.
Brown, F. B., and others. Discussion on decay in timber. Trans. Canad. Soc.
Civ. Engin. 29, pt. 1 : 324r-365, 13 pi. 1915.
Bibliography of timber destroying fungi, p. 339-340.
Clinton, George Perkins. Report of the botanist for 1915. Connecticut Agr. Expt.
Sta. 1916, pt. 6: 421-487, pi. 17-26. August, 1916.
Notes on plant diseases of Connecticut; diseases of plants caused by nema-
todes; powdery scab of potatoes; potato spraying experiments, 3d report.
Literature, p. 461-462, 469.
Culpepper, Charles £., Foster, Arthur C, and Caldwell, Joseph S. Some effects
of the blackrot fungus, Sphaeropsis malorum, upon the chemical composition
of the apple. Jour. Agr. Research 7, no. 1 : 17-40. October 2, 1916.
Literature cited, p. 39-40.
Fromme, Fred Benton. Facultative heteroecism (r) of Peridermium harknessii
and Cronartium quercus. Phytopathology 6, no. 5: 411-412. October, 1916.
Gorkum, Nicolas van. A molestia do olho da canna, ponto- de vegetaySo, corag&o
e peciolo da bandeira. Bol. Min. Agr. Indus, e Com. [Brazil] 4, no. 2: 105-112,
3 col. pi. Abril/Junho, 1915.
Gtissow, Hans Theodor. The grain rust in the prairie provinces. Agr. Gaz. Can-
ada 3, no. 10: 861-864. October, 1916.
Hawkins, Lon Adrian. Growth of parasitic fungi on concentrated solutions. Jour.
Agr. Research 7, no. 5: 255-260. October 30, 1916.
Literature cited, p. 259-260.
Hedgcock, George Grant, and Hunt, N. Rez. Dothichiza populea in the United
States. Mycologia 8, no. 6: 300-308, pi. 194-195. November, 1916.
Literature cited, p. 308.
Hesler, Lexemuel Ray. Black rot, leaf spot, and canker of pomaceous fruits.
New York Cornell Agr. Expt. Sta. Bui. 379: 49-148, fig. 18-37, pi. 7-14. 1916.
Bibliography [annotated], p. 126-148.
Home, William Titus. The importance and prevention of wood decay in fruit
trees. Mo. Bui. State Com. Hort. [California] 6, no. 8: 278-282, fig. 95-96. Au-
gust, 1916.
1 This list aims to include the publications of North and South America, the West India Islands, and
islanHs controlled by the United States, and articles by American writers appearing in foreisn Journals.
All authors are urged to co5perate in making the list complete by sending their separates and by mak-
ing oorrertions and additions, and especially by calling attention to meritorious articles published outside
of regular journals. Reprints or correspondence should be addressed to Miss E. R. Oberly, librarian.
Bureau of Plant Industry, U. 8. Dept. Agric, Wsshington, D. C.
80 Phytopathology [Vol. 7
Hotaon John WiUUm. The longevity of Bacillus amylovorus under field conditions.
Phytopathology 6, no. 5: 40(M08, 4 fig. October, 1916.
Johnson, James. Enfermedades del tabaco y manera de combatirlas. Pt. II-IV.
Hacienda IS, no. 1: 26-28, illus., Octubre; no. 2: 6^-64, illus., Noviembre; no.
3: 01-03, illus., Diciombre. 1016.
IH. 1 noted in previous list.
Host plants of Thielavia basicoia. Jour. Agr. Research 7, no. 6: 28^-^00.
pi. 18-10. November 6, 1016.
Johnston, John Robert. Phytopathological work in the tropics. Phytopathology
6, no. 5: 381-386. October, 1016.
Jones, Lewis Ralph, Johnson, Aaron Guy, and Reddy, C. S. Bacterial blights of
barley and certain other cereals. Science n. s. 44, no. 1134: 432-433. Septem-
ber 22, 1016.
Long, William Henry. The aecial stage of Coleosporium ribicola. Mycologia 8,
no. 6: 300 311. November. 1016.
Peridcrmxum rihiciAa.
Matz, J. A meth<Mi to induce HiM>ruIation in cultures of Botryosphisria beren-
geriana. Phytopathology 6, no. 5: 387 380, 1 fig. October. 1016.
Meinecke, Emillo Pepe Michael. I^h vanillidres dc Talnti &, de M(M)rea. Rap-
port pr^>nt^* tt Mr le (louvcrneur des cstablissemcnts fran^ais de TOc^anie.
& a MM. Icfl niemhrcH de la chambro d'agriculture. 44 p. Papeete, 1016.
Maladies it leurti rauM'M, p. 13-28; suppression des causes des maladies de
la vanille, p. 20 32.
If elchers, Leo Edward. Diseases affecting Sudan grass. Kansas Agr. Expt. 8ta.
Bui. 212: 16^-10, pi. 4-^5. 1016.
Kernel smut; treatment of s4M>d to kill smut; bliglit; seedling root disease.
Plant disease's afferting alfalfa. Upt. Kannas State Bd. Agr. 85, no. 138:
330 ;i53. fig. 2S2 a>3. 1010.
Vromycrn atriatuB Schrcet; IWutloprziza mediraginin (Lib.) Sacc; PhyUo$ticta
sp; PUoaphfrrulina brionuina Pollarri; Peronospora trifoliorum De By; /?Ai-
loelimia violarta Tul; PnewiomonaB mniicafjinis; yellow top; stem cracking
{Phnma Hp. K
Krrata note: V\H\n statement of the author, the geniu) Phomo|)sis, p. 330
and .'MO, Hhould reiul Phoma.
Melhus, Inring E., Rosenbaum, Joseph, and Schultz, Eugene S. S|K)ngos|N>ra
Hubtorranea and l'h(»ma tulK*n»Ha on the Irinh potato. Jour. .\gr. Research
7, nf». :>: 213 254, 1 fig., pi. A. 7 14. OctolMT .«). 1016.
Ifash, George Valentine. Injury to evergrcenH. .Jour. New York Bot. Card. 17,
nt>. JSri: 170 lH."i. OrtolnT. 1016.
Ifowell, WlUlam. The dying of citrus tn^en: :i coinpariHon. .Agr. News [Barbaiios]
15, no. 370: 'M\k\'MV:. N«»v<iii»mt 4. 1016.
O'Gara, Patrick Joseph. Orcurrcnre of yellow leaf rust of wheat (Puccinia gluma-
runi» in thi* Salt Lakr valley, I'tah. Science n.H. 44, no. 1130: 610-^11. Octo-
Imt 27. 1016.
Pammel, Louis Hermann, King, Charlotte M., and Seal, J. L. Studies on a Fusarium
diM*as4* of corn and sorglium. il'reliminary. j Iowa .Agr. Kxpt. Sta. Research
Bui :»: 113 VM\. \:^ fig 1016.
Litcrattirr of corn ri»tM. p. 11') IIH.
Reimer, Frank Charles. .\ promiiting new |>i*ar ntock. .Mo. Bui. State Com. Ilort.
(California) 6, no. :>: \m 171, fig. oO 60. .May, 1016.
Pyrua calUryana. Very resistant to blight.
1917] Literature on American Plant Diseases 81
Rumbold, Caroline. Pathological anatomy of the injected trunks of chestnut trees.
Proc. Amer. Phil. Soc. 66, no. 6: 48^-493, pi. 15-18. July, 1916.
Literature cited, p. 493.
Schneider, Albert. A parasitic saccharomycete of the tomato. Phytopathology
6, no. 5: 39^399, 4 fig. October, 1916.
Shear, Cornelius Lott. False blossom of the cultivated cranberry. U. S. Dept.
Agr. Bui. 444, 7 p., 4 pi. 1916.
Literature cited, p. 7.
Cause not known.
Smith, Clayton Orville. Crown gall or plant cancer. Mo. Bui. State Com. Hort.
[California] 6, no. 6: 201-211, fig. 71-72. 1916.
Smith, Erwin Frink. Tumors in plants. Science n.s. 44, no. 1139: 611-^12. Octo-
ber 27, 1916.
Stewart, Fred Carlton. Observations on some degenerate strains of potatoes.
New York State Agr. Expt. Sta. Bui. 422: 310-357, 12 pi. 1916.
Stoddard, £. M., and Moss, A. £. Cutting out chestnut blighted timber. Connecti-
cut Agr. Expt. Sta. Ann. Rpt. 1916, pt. 6: 488-496, pi. 27-28. August, 1916.
Stone, George Edward. Injury to vegetation resulting from climatic conditions.
Jour. New York Bot. Card. 17, no. 202: 173-179. October, 1916.
Thorn, Charles, and Currie, James N. Aspergillus niger group. Jour. Agr. Re-
search 7, no. 1: 1-15. October 2, 1916.
Bibliographical footnotes.
Tisdale, W. H. A Melanconium parasitic on the potato. Phytopathology 6, no. 5:
390-394, 3 fig. October, 1916.
Relation of soil temperature to infection of flax by Fusarium lini. Phyto-
pathology 6, no. 5: 412-413. October, 1916.
U. S. Department of the Agriculture. Bureau of Plant Industry. Report of the
chief, [19151/16. 18 p. 1916.
Plant pathological investigations, p. 3-6.
Federal Horticultural Board. Report [1915]/16. 14 p. 1916.
Service and regulatory announcements. August: 93-101, September
30; September: 103-129, 1 fig., November 16. 1916.
Weir, James Robert. Phacidium infestans on western conifers. Phytopathology
6, no. 5: 413-414. October, 1916.
Pinus ponderosa and P. jeff'reyi, hosts for Razoumofskya americana. Phy-
topathology 6, no. 5: 484. October, 1916.
Whetzel, Herbert Hice, Hesler, Lexemuel Ray, Gregory, Charles Truman, and Ran-
kin, William Howard. Laboratory outlines in plant pathology. 207 p. Ithaca,
New York, 1916.
[Phytopathology, for December, 1916 (6: 419-454) was issued Novem-
ber 29, 1916.]
PHYTOPATHOLOGY
VOLUME VII NUMBER 2
APRIL, 1917
THE PERFECT STAGE OF GLCEOSPORIUM VENETUM
Walter PI . B u k k h o l d e it
With Three Fkjures in the Text
During the oarly siuniner of 1914 while studying the anthracnose
disease of the raspberry at Brant, New York, a peculiar ascomycete was
observed by the writer. The fungus although not of general occurrence
was found only in the anthracnose lesions (fig. 1), and arose from the
stroma of the pathogene GloBOsporium venetum Speg. This led to the
belief that there was a possible connection between the two fungous
fonns and a niunber of inoculation experiments were conducted in order
to verify this assumption.
It was difficult to obtain ascospores for making inoculations owing to
the scarcity of the ascocarps. Furthermore the ascospores w^ere borne in
the same lesion with the conidia of Glceosporium venetum and a separation of
the two types of spores was practically impossible. It was finally decided
to use spores from a culture of the fungus developed from a single
ascospore.
Several attempts were made to isolate the fungus. The poured plate
method first employed was discarded later on account of the difficulty
in obtaining ascospores sufficiently removed from the conidia, the latter
usually being in great abundance. A second method and similar to one
used by Barber^ was also tried. This consisted in crushing the ascocarps
in a drop of sterilized water on a sterilized slide. A glass tube with a
bore of about 3 mm. was drawn to a capillary tip at one end; to the op-
posite end was fastened a piece of rubber tubing about 40 cm. in length.
The free end of the rubber tube was placed in the mouth and by manip-
ulating the glass point with the hand, spores could be drawn into the bore
of the tube. The great difficulty in using this method with the fungus
under consideration was the fact that the ascospores were very gelatinous
and had a tendency to adhere to the glass slides, refusing to enter the
* Barber, M. A. On heredity in certain microorganisms. Kansas Sci. Bui. 4:
3-48. 1907.
1917] Burkholder: Plectodiscella veneta 85
globules and pigments which give color to the fungus. Only on rare
occasions and on media containing a small percentage of agar do fila-
mentous hyphae extend for any distance from the sclerotia-like formations.
This growth on artificial media although identical with that of (?. venetum
is decidedly different from that of any species of Gloeosporiimi which has
a perfect stage belonging to the genus Glomerella.
As the writer has continually foimd to be the case with cultmres of G.
venetum, difficulty was encoimtered in finding conditions favorable for
the sporulation of the fimgus. It was finally observed, however, that a
sudden change in the humidity of the cultmre tube caused a production
of conidia which were obtainable in sufficient numbers for use. In order
to effect this change the fungus was grown on three-per-cent potato agar
until large sclerotia-like masses were formed. These masses were trans-
ferred to sterilized bean pods in tubes which contained several centi-
meters of water. The cultures were tiien incubated at a temperatmre of
24°C. and at the end of three days numerous conidia were produced which
were identical with the conidia produced in cultmre by G. venetum. It
was also observed that this sporulation was not continuous, but ceased
after the first production of spores. Fiu-thermore, a cultiu*e of the fxmgus
subjected from the beginning to a moist condition produced no spores or
at least, but relatively few. This apparently indicates that the sudden
increase in himiidity acts as a stimulus to spore formation.
By dropping these fimgous masses bearing conidia into a small amount
of water the spores readily fall off and can be sprayed over the infection
court. The germination of these conidia is fairly rapid, but the percent-
age of germination is low. In most cases not more than five or ten per
cent of the spores germinate.
INOCULATION EXPERIMENTS
Early in the winter of 1914 a nxmiber of roots of the Colxmibian variety
of the raspberry were obtained and planted in the greenhouse. Owing
to the earliness of the dormant period and to the unfavorable conditions
arising within the greenhouse, the plants grew slowly, and gave a very
stxmted growth. All inoculation experiments with these plants gave
negative results. The anthracnose lesions appear only on tender suc-
culent canes and apparently the canes which had developed slowly on
the greenhouse plants were too hard for the fungus to infect.
Later, about the first of March, 1915, a few raspberry plants of a red
variety were secured which were tender and growing rapidly. On March
4, two canes were sprayed with a suspension of conidia from a culture of
the fungus developed from a single ascospore and the canes were covered
86 Phytopathology [Vou 7
with bell-glasses lined with moist filter paper. These glasses were plugged
at the top with cotton and allowed to remain over the canes for two days
before removing, while two other canes in the same bed remained imtreated.
On March 20, small purple spots had appeared on one of the canes. These
infections grew slowly, much slower than an anthracnose lesion develops
under field conditions, but spots typical of those caused by Gkto^porium
venetum were produced. Miscroscopical examination of the spots showed
conidia of G, veneium.
Again on April 15, four very tender canes of a black-cap variety of
raspberry were sprayed with a suspension of conidia as above. Bell
glasses were placed over the plants as in the previous experiments and one
check plant was tised. A sample of the conidia used was placed in a drop
of water on a slide, and about eight per cent of the spores germinated.
After one week, April 21, a nimiber of small purple spots had appeared on
the four canes, and these later developed into t>'pical anthracnose lesions.
The check plant remained healthy.
THE DEVELOPMENT OF THE A8CIGEROL8 STAGE
From the positive results of the inoculation experiments and also from
the examination of the fimgus in culture it is evident that the ascomycete
under consideration is the perfect st-age of Gkeoftporium venetum. The
8>'stematic position of the fungus, however, is rather diflficult to determine.
Its morj)holog>' is entirely different from the perfect stage of any species
of (ilcDosporium previously described. This, however, is not siuprising
as (7. veneium has always l)ec»n considenMl distinct from the other species
of that genus.
The ascig(»n)us stage of the fungus wjis first ol)served on the hybrid
raspl)err>' commonly known as Kubus negleclwt. I^ter it was collecte<l
in various |wirts of Ne^* York Stati* on the bbick rasplx»rr>' (Rubus occi-
dentalis) and the American red rasplxTry (Kubus idceus var. aculeaiwimuB) .
RcHis' alMo rei)orts having found it iti Washington on the blackl)err>'
(Rubua sp.).
During the siunmer following the* discovery of the ascigerous stage
clos4» ol>ser\'ation was kept of the fungus on th(» young canei* to detennine
when the Jisconiqw first lK»gan to <lcv(»lop. This pn)ve<l to be about the
middle of August. At this time the fruiting Inxlies which greatly resemble
those in the family Myriangiaceie apfK^iir as minute s|K)ts, deep brown to
black, singly or in grou|>s s<'attenHl over the huff-colored and sunken por-
tion of the anthracnose* lesion. These spots an» Imrely visible to the €>•«
* Rm"^. H. L. Kx|N*nmental (ipra>nnK for hlarkbeiT>' anthracnose in 1915. Watt-
em WanhinKton Kxp. Sta. Mo. Bui. 3*: 1-10. 1915.
1917] Burkholdbb: Plectodiscella veneta 87
and only so on account of the contrast in color with the surrounding tissue.
After passing the winter the entire lesion assumes a dark brown color and
then the pustules are observed with great difficulty even with a hand lens.
A careful examination of the diseased area upon which the asco arps
are found proves it to be a typical anthracnose lesion. The buflf-colored
portion is fimgous tissue, more or less plectenchymatous in structiu'e.
It is composed of very small hyphae which are difficult to distinguish
imless carefully stained and it is the same tissue from which arise the
conidiophores of the GlcEOsporixun stage. The ascocarps arise from the
stroma and are pulvinate structures usually circular in outline, but they
frequently coalesce, forming spots of various shapes. They are approx-
imately 0.07 by 0.07 to 0.37 mm. in diameter. The tissue of the ascocarp
is more or less pseudoparenchymatous with larger and thinner-walled
cells than those of the stromataceous tissue (fig. 2). The outer layer of
the ascocarp is composed of thick-walled brown cells which form a shield-
shaped structiu'e less perfect, however, than those observed in the family
Microtheriaceffi. When the fruiting body is matiu'e the cells of this outer
layer split apart- in a stellate manner and crumble away. Within the
shield the ascocarps are hyaline and contain the asci which are scattered
irregularly through the fungous tissue. There is no differentiated cavity
for the asci.
The asci were first observed in the immature condition at which time
they appeared as globose bodies containing a homogenous mass of proto-
plasm, and greatly resembling thick-walled oogonia. These asci may lie
in contact with each other but frequently they are separated by the fim-
gous tissue. The mature ascus is thick-walled and measures 24 to 30/4
in diameter. In a few cases the ascus has appeared to be slightly stalked
and attached to the base of the cavity in which it is borne.
In the autmnn or more often in the spring the asci matiu-e and the
homogenous mass of protoplasm gives place to eight four-celled ascospores.
These spores which are borne parallel to each other in the ascus, are hyaline
with gelatinous walls, and constricted at the septa. The basal cell is some-
what more obtuse than the apical cell. The mature ascospore measiu'es
18 to 21/i in length by 6.5 to 8/u in diameter.
In the formation of these spores the middle septum is laid down much
earlier than the other two, and for this reason it is not imcommon to find
two-celled spores. With the division of these cells giving rise to the
foiu'-celled condition, the constrictions are not so great as at the first
septimi. Occasionally one of the cells fails to divide and an ascospore of
three cells is formed. During the formation of the spore the disintegration
of the fimgous tissue about the asci takes place and with the rupturing
of the outer layer of the ascocarp the asci are exposed. Frequently the
Phttopatholoot
[Vol.7
asci lying in this exposed condition surrounded by the remainder of Um
•scocarp give the appearance of a true discomycete. This, homrer, is
duo to the persistency of the outer cells of the ^eld-like layer whidi
Pia. 2. «Crou-«iction or AscocAxr or PLCcrootacsLLA vbmbta
Vuioua itKces ia th« dcrelopmeat of the uci u« ■hown
1917] Burkholder: Plectodiscella veneta 89
covers the immature ascocarps. With the presence of sufficient moistiu-e
the exposed asci elongate approximately three times their usual length.
This process is very rapid and may be observed imder a microscope when
a fragment of tissue containing asci is placed in a drop of water. The
lower portion of the ascus remains fastened in the cavity in which it was
borne, giving a conical shape to the body which raises itself above the sur-
rounding tissue. The spores gather at the tip of the ascus and from there
are ejected into the air. They have been caught above the lesions at
a distance of one centimeter.
In a single ascocarp all the asci are never in the same state of maturity
and the ascocarps also seem to vary in this respect. Mature ascospores
were first observed about the first of Jime while inunatiu'e spores were
present in August.
GERMINATION OF SPORES
When placed in tap, rain or distilled water, or on nutrient agar, the
mature ascospores germinate readily (fig. 3). They swell somewhat and
within less than two hours a short sterigma is produced from one or each
of the cells. A sprout conidiimi is formed which is oblong to elliptical
and identical with the conidia of the fungus. When fuUy mature the
sprout conidia drop from the sterigmata but do not germinate immedi-
ately. After a short period of rest, twelve to twenty-four hours, a germ-
tube is sent forth and mycelium is formed. When an ascus is placed in
a drop of water or on agar the spores within TviU germinate by sending the
sterigmata through the wall of the ascus and produce the sprout conidia
on the outside. These, in turn, germinate. After Ihe production of
the secondary spores, however, the ascospores shrivel and disintegrate.
SYSTEMATIC POSITION
The morphology of the fungus of the raspberry anthracnose, especially
the character of the asci scattered irregularly through a pseudoparenchyma
is similar to that of the old family Myriangiacese. In a revision of this
family some years ago by von Hohnel* but five genera out of twenty-three
were retained, and since then but one new genus, Ascostralum Sydow^ has
been added. The perfect stage of Glceosporium venetum, however, does
not appear to fall in any of these genera, nor in any of the genera of closely
' Hohnel, F. von. Fragments zur mycologie VI. Mitt. 244. Sitzungsber. M-N
Classe, k. k. Akad. Wiss. Wien. U8: 349-376. 1909.
* Sydow, H. von and Sydow P. von. Beschreibungen neuen sUdafrikanischen Pilze.
Ann. Myc. 10: 41-42. 1912.
90 Phytopathology [Vol. 7
related families. More recently Woronichin'^ described a new genus,
Plectodiscella, based on a single species which he found occurring on the
leaves of the apple and pear. This genus is closely related to ELdnce of
Raciborski* butdififers mainly in that the stroma is not borne beneaUi the
epidermis. Plectodiscella Piri, the representative of the genus, is so simi-
lar in morphology to the ascigerous stage of GUeosporium venetum Speg.
that apparently there is a distinct relation between the two. A brief
description of Woronichin's species is here set forth: A more or less im-
perfect stroma is formed in the epidennal and sub-epidermal cells of the
leaf, which is at first sub-cuticular. From this arises a fimgous tissue in
which are borne irregular globose asci, each containing eight four-celled asco-
pores. In some instances the asci are separated by the fimgous tissue
and in others they lie in contact with each other. Woronichin is uncer-
tain as to the nature of this tissue l>etween the asci. He says, **Wa«
fiir Elementc die z^'ischen den Ascen l)efindlichen Zwischenraume ausfQl-
len, gelang es nicht genau aufzuklaren." He does not consider, however,
that it is cellular. This is also the first impression received in regard to
the raspberr>' fungus, due to the fact that the cells are minute and disinte-
grate very ejirly. The cellular strticture of the fungus on raspberr>'
was determined only on young material and then after it was fixed and
8taine<i. In Plectodiscella Piri a shield-like arrangement composed of one
layer of dark cells covers each ascocarp and later breaks apart in order
that the asci may be ejcposed. In the perfect stage of Glaoiporium vene-
turn this is present but is clearly seen only in the inmiature stages before
rupturing occurs. Woronichin does not refer to an imperfect stage for
his fungus.
Plectodiscella Piri is considered to be far enough removed from the
MyriangiacesB or any of its closely related families to be placed in a new
family. Tliis, Woronichin describes as Plectodiscelle® and states that
it occupies a s>^tematic position somewhere lx»tween the Plectascales
and the true Discomycetes. Here he places his fungus P. Pin* but gives
no exact characters for his genus. Only the family and species are
descril)ed.
Taking all characters into consideration, the pcTfcH't stage of GUto-
sporium venetum Speg. appears to belong to this genus and therefore the
following name is proponed:
• Woronichin, N. M. Plertodiscella Piri, diT Vcrtrctcr einorneuen ascomyc«teii
Gnippe Nfycol. Centralh. 4: 225-233. 1914.
* Harikxirfiki. M. Kluiomv Har. nov. f(en. Parasitiiiche Algen und Pilte Jara't
1: 15-16. IWO.
1917] Burkholder: Plectodiscella veneta -91
Plectodiscella veneta sp. nov.
Stromattbns solitariis vel gregariis, pvlvinaiiSy epidermide fusca diacoida,
mox dehiscerUe, intus contextu hyalinOf pseudoparenchymatico vel indistincto,
plerumque pluriloculigeris, loculis monasciSy irregulariter sparsis; dscis
globosis, 8 sporis, 24rS0n; sporidiis ovoideo^llipsoideis, saepe flexis, hyalintOy
SseptatiSy canstrictis, ceUuLa basilare cbtusay 18-21 x 6,6 x 8fi.
Hob, In ramis caulibuaque vivis Rvbi occiderUalis, R. idaei var.
aculeatissiini et R, neglecti. New Yorky America bcreale. Status coni-
diaphorus eat Glceosporium venetum Speg,
Cornell University
Ithaca, New York
PUCCINIA SUBNITENS AND ITS AECIAL HOSTS
Ellsworth Bkthkl
Puccinia subnUens Diet. \b a common rust on Distichlis 9pieaia from
the Atlantic to the Pacific coast. The telial host is especially abundant
in the alkaline soils of the desert regions of the western United States.
Prior to 1904, Chenopodium album was the only known aecial host of this
rust. In the summer of 1904, Rev. J. M. Bates, from field observations
in Nebraska, concluded that aecia on species of Cleome, Sophia, Lepid-
ium, Erysinuim, and Salsola were related to Pxiccinia subnilens. These
suggestions iiiTre conmiunicated to Dr. J. C. Arthur, who later in the
summer succeeded in growing the teliospores on these hosts, thus con-
firming the deductions made by Bates. Dr. Arthur, in giving the results
of these cultim»s, remarks (Joiu*. Myc. 11: 50-67), *'\Ve have here a dem-
onstration of the remarkable fact, not known for any other species of
rust, that Puccinia subniiens has aecia growing \iith equal vigor on three
families of plants." I-.ater, Arthur grew it on Capsella (Bursa) sp.,
AtripUz hastataf and doubtfully on Sarcobaius sp. He has grown this
species on nine or ten genera in three families.
For ten >'ears or nrort*, the writer has observed the aecia of this rust
abundant in Colorado on species of Polygonum, (chenopodium, Lepidium,
Capsella, Cleome, Salsola, and Abronia, and in 1912 made successful cul-
tures on all of these genera except Abronia. During the past summer,
aecia which seemed immistakably related to Puccinia subnitens were
foimd on plants of several other genera, and cultures were made to de-
tennine the correctness of these observations. Likewise all previous
cultun»s i*-ere repeated with the result that the aecia were produced on
22 s|MM*ies in 6 families, and 15 genera. The following is a list of suc-
cessful rtiltures.
1. Polygon ACK.*:: Polygonum aviculare L., P. erecium L., P. ramonstt-
mum Mirhx.
2. ('iiknoi»<>i>ia<k.k: SaUola peMiftr \. Nc»L*<., Chenopodium album L.,
r. ijhiHcxim L.. r. Uiiiceolatum Muhl., (\ pagonum Reich., Manoltpi^
nutUillittnn iK. iV Sj (Sreene, Kttchia scojmria (L.) Roth.
3. .\MAliANTHArK.K: Awaranthu^i rttroflexus L., A, blitoides S. Wats.
4. Ny( TA<flNArK.K: Ahnnn^i fnigrans S\i\{.
5. Ckitikkk.*:: ('apsrlln liursa-iHistoris Me<lik.. Lepidium dmsiflorum
S<'hnid., L. medium (ireene. Erysimum oHperum DC., Sophia pinnaia
1917] Bethel: Aecial Hosts of Puccini a Subnitens 93
(Walt.) Britt., Roripa palustris (L.) Bess., Thlaspi arvense L., Sisym-
brium aUisaimum L.
6. CAPPARiDACEiB: Cleomc semUata Pursh.
It will be observed that the above six families constitute two groups.
The first four comprise a group of contiguous families, and the last two
another group, likewise contiguous but rather remote from the first.
Mcidium fumaricLcearum Kell. & Swingle on Corydalis is probably ne-
lated to P. subnitens, though no cultures were made. If this connection
should be established it would add another family, Papaveracese, con-
tiguous with the second group, a total of seven famiUes.
It is not uncommon to find the aecia in abimdance on a half dozen or
more host plants at one place, and it manifests only slight racial tenden7
cies, though it seems to infect certain hosts, such as Thlaspi, Kochia,
Monolepis, Amaranthus, Roripa, and Erysimum very sparingly. The
aecia occm* in abimdance on plants of all other genera Usted above.
Stanleya pinnata (Pursh.) Britt. bears a large orange-red aecium char-
acteristic of P. subnitensy however, this host failed to become infected,
though seven cultures were made under the same conditions, and with
the same material that was used in the successful cultures on other
hosts. Cultures were attempted on Atriplez hastata L., A. canescens
James, A. confertifolia S. Wats., and Sarcobatus vermiculatus (Hook.)
Torr. with no results. Arthur has reported successful cultures on Atrip-
lex hcistata L. with teUospores from Delaware, and on Sarcobatus vermi-
ciUatiLs (Hook.) Torr. with teliospores from Nevada. The aecia on the
latter host, which closely resemble those of P. subnitens, are related
chiefly, or entirely, at least in Colorado, to Pucdnia Ixixuriosa Syd. on
Sporobolus airoides Torr., as has been shown by the writer by several
successful cultures the past season, both from aeciospores and teUospores,
so that there can be no doubt of this relationship. Further, many sow-
ings of teUospores of P. luxuriosa on the aecial hosts of P. subnitens made
through two seasons gave negative results.
Mcidium Abronioe E. & E. was described on Abronia sp. from Fort
Collins, Colo. Many cultures both in the field and the garden show
that it is the aecial stage of P. subnitens. It is common on Abronia
fragrans Nutt. in Colorado and occasionaUy collected on A. eUiptica A.
Nels., and A. micrantha A. Gray.
Late in the summer cultm'es were attempted on Beta, BUtum, and
Portulaca but no infection resulted, presumably for the reason that
the teUospores' had probably already germinated. Successful cultures
were obtained on either Raphanus, or Brassica, but the plants died be-
fore developing sufficiently for determination. The aecium on Cleo-
mella, as noted by Arthur, also is probably related to P. subnitens. Cul-
94 Phytopathology [Vol, 7
lures vnW lie made again next Heason on Stanleya, Atriplex, Corydalis,
ClmmicUa, Beta and plants of some other suspected i^encra, and if these
prf)vc to be aecial hosts of this rust, which seems very probable, we shall
have a grand total of more than a score of genera in seven families-
remarkably large nimil)er of aecial hosts for a single species of rust.
Colorado State Museum
Denver, Colorado
CONTRIBUTIONS TO OUR KNOWLEDGE OF THE WHITE
PINE BLISTER RUST
W. A. MCCUBBIN
I. MODE OF INFECTION ON THE PINE
Only indefinite references to the method of infection of the pine by Cro-
nartium ribicola have appeared in current literature. From these refer-
ences one gathers the impression that infection takes place through the
bark, and probably by way of woimds or abrasions. Having an oppor-
timity for studying a considerable nimiber of pine infections in 1916,
some attention was given to this point, and records were made of the
origins of cankers where such origins could be determined.
In most cases the determination was not difficult, owing to the fact
that in a healthy pine branch the fimgus spreads out from the court of
entry in a very regular and equal manner, and that its progress is marked
by swelling or discoloration or both, or else the cortical tissue is killed in
an equally radial fashion. By taking note of this habit one can readily
locate the point of 'original infection in most cases, especially in the
earlier stages.
TABLE 1
Records of specific cases to show mode of infection of pines by Cronartium ribicola
LOCALITY *
Secords
Four-mile Creek
Cookstown
Totals
I>rUMBEH OP
INPBCT10.V8
177
38
792
1007
ORIGIN' OF LBfllON
IjesJ fasdcleB
148
M
743
925
Wounds
8
1
5
14
Undetermined
21
3
44
68
Very early in this study it became apparent that the chief mode of
infection was by way of leaf fascicles through the so-called short shoots.
In these pines, which were all healthy and which grew in situations where
they were fairly free from accidents, wound infection played but a very
small part..
According to the tabulated results about 92 per cent of these young
bUster cankers originate in leaf-bundle infection. This percentage in-
96 Phytopathology [Vol. 7
eludes only those cases where the point of ori|^n could be confidently
established, but it is highly probable that a large proportion of the num-
ber listed as undetermined should also find a place here, and it might
not be overstepping the mark to ascribe at least 95 per cent of thede
blister cankers to leaf fascicle infection.
One may consider that the sporidia from the currant leaves are lodged
among the bases of the needles and from this position can then attack the
short shoot which bears these leaves. In a number of instances a few of
the leaves on such shoots were found to be dead while the rest of those in
the fascicle were quite healthy; in other cases all the leaves in the fascicle
had l)een destro>'ed and often the short shoot and even a small area in
the cortex at its base were also killed. In milder cases, especially where
the growth of the tree was very vigorous, the fungus did not kill either
the leaves or the short shoot, but induced in the latter a pronounced
stimulation of growth, so that the short shoot became enlarged and
buU)ous in appearance.
In these peculiarities of short shoot infection may lie a possible expla-
nation of the year of dormancy which so evidently obtains in a great ma-
jority of C4u*es. If. during the summer after infection, the fungus pro-
gresses only into the short shoot or slightly beyond it into the adjacent
cortex, it would l)e difficult to recognize these minute symptoms and
then» would \h* the so-called dormant year.
II. LIFE CYCLE OF THE FUNGUS ON THE PINE
In general the tendency has been to regard the life of Cronartium rtW-
coln on its pine host as more or less indefinite, varying from one to two
y<»ar8 up to six years or more; that is, from the time of infection \mtil
aeria are pnxluced from one to six years might elapse. In the study of
this disea^ in Ontario in 1916 evidence has come to hand which indicates
that the fungus tends to reach the aecial stage in a fairly definite period,
but tliat this noniial course of development may l)e shortened or length-
onetl lMH»ause of c<»rtain favorable or unfavorable factors.
In the Niagara Penin.»<ula in 1916 there were found a numl)er of young
pint* infections on tre<»s growing close* to black currant-s, and though these
currants have Innnx badly nisted since and incluchng 1914, there is reason
to think tliat no nL»»t was present on them Ix^fore that season. In any
CUM* no >«ign of infection was visible on these pines in 1915, although they
wen* ciTtainly ex]x)sed to infection during the previous year. Moreover,
in the summer of 1916 no (hs4»as<» was met with on the groH'th of 1915
although the.*^» pines must have n»c(»iv(»(l infection in 1915. On the other
hand, th<»S4» five lots of pines dcvelo|MMi 22.3 infections in 1916, all on the
1917] McCubbin: White Pine Blister Rust 97
growth of 1913 and 1914. It seems reasonable to think, therefore, that
during the year after infection there are no symptoms of a visible natxire
in infected pine branches. Additional evidence on this point recently has
been obtained from another district (Cookstown, Simcoe County, On-
tario), where a young nursery row of white pines was severely infected
from black currants growing side by side with them. Out of the 1412
blister cankers recorded from these rows not one was found on the growth
of 1915 although the pines were certainly subject to infection in the
previous year. On the twig growth of 1914 there occurred some 286
cases, which number gives a strong indication as to the yearly infection
that might be expected here.
While the above evidence from these two cases is scarcely conclusive,
it is sufficiently extensive tad clear-cut to warrant the assumption that
in the great majority of cases the season following infection is a "dor-
mant" year.
When the character of all the 509 infections recorded above is examined
further another point becomes clear. With one exception all of these
cankers were in what might be termed the swelling stage, the cortex being
typically swollen into a spindle and usually discolored. If these cankers
can be taken to represent the normal course of the disease, and since they
include all the infections foimd on the 1914 twigs of these quite normal
trees there seems to be no reason why they should not be so considered, —
then the third season of the diseare is apparently characterized by the
appearance of the first visible symptoms, the swellings just mentioned.
And if the formation of aecia from these swellings be assumed to take
place during succeeding years a fairly normal life cycle will have been
obtained. Summarized it would run thus: first season, infection in summer
and autumn; second season, dormant period; third season, swelling stage;
fourth season, aecia. There is evidence, however, that in the majority
of cases the swelling stage may last for two years before the production of
aecia. This evidence has been obtained partly from the Cookstown case
already mentioned and partly from the Secord case where 177 swellings
were found in 1916 on yoimg pines growing close to black currants.
When these 177 swellings were arranged in a curve representing the
number of them that had occurred on each yearns growth it was found that
the apex of the curve came in 1914; that is, there were more infections
on the growth made in 1914 than on that of any other year. WTien a
curve was prepared similarly from the data obtained at Cookstown, it
was of another type, having the largest number of swellings on the growth
of 1913 (fig. 1). The nature of this curve thus suggests very strongly
that the swelling stage may ordinarily last two years before the aecia are
produced.
98 Phttopatholoot (Vol. 7
It is to be noted that the Cookstown curve repreaenta infectiotu begun
in several successive yean vhile the Secord curve contains infections of
only one year, that of 1914. An attempt was made to compare the two
wries of results on a basis of something like equality by supposing that
in addition to the infections started in 1914 in the Secord case a similar
series of infections had begun in 1913. Assuming that the mrellings
which would presumably arise from these earlier infections in 1915,
would still remain in the same stage during 1916, a curve was then con-
Kor expluiation hc text
Hlnirtt^d which would iiicluili' thif> h\-iMtthetiiial xeriem of cankers along
with those sc-tually pn^etit. Surh a cnrw would contain the infections
of two uur«*iwiv(* ypiirf and would ihuo n-wniblo ver>- closely the Cooks-
town i-urvc. When the ntmilxTs in thi!< n-conmnicted curve were doubled
in order 1o obtuiii ii In-ttcr roin|K>ri.-«m with the Cookstown curve, it was
Bc«>n thsit fxcppt for minor irreKularities the two ('iir\'C!' are practically
iilcnticul.
The Won*- coiicordiuici: of this sup)x>«'d ciiw with the results of the
Hctiud :tiir\'py prrtvidc:* n xtrikinK confinimtion of the indication already
1917]
McCubbin: White Pine Blister Rust
99
given in the Cookstown curve of the continuance of the swelling stage
for two years.
The evidence may be presented in another way, by following the course
of the disease year by year in a number of shoots of 1911, exposed as in
the Cookstown case to a more or less constant annual infection from cur-
rants. The number of infections started in these pine twigs in 1911 may
be represented by X, those started in 1912 by 7, and those in 1913 byZ.
Later infections may no doubt take place in these shoots, but smce the
number of such infections is known from other considerations to be very
small, they may be neglected for the purpose in view. A small letter
table 2
Probable development of the blister rust in pine branches based on a four-years eyeU
TSAR
TBARLT PBOOBE8S OW
NXTMBSR OF 8WKLLIN08
ESTIlf ATBD PBOPOB-
TIONB BASED ON BBCOBD
COOKSTOWN SUBYBT
DISBA8B
PBRSENT BACH TXAB
CA8B
rXGUBBS
1911
X*
0
0
0
1912
X^-fY*
0
0
0
1913
x'+yd-hz*
X
300
286
1914
XHY»+Z<*
Y
100
390
1915
Xb_|.Y»>+z«
Z
10
83
1916
X*>+Y*>+Z»>
0
0
22
TABLE 3
Probable development of white pine blister rust in pine branc?ies based on a five^ffears
cycle
TBAB
TBABLT PBOOBX88 OP
DBUASB
NDMBBB OP SWBLLINaS
PBBSBNT BACH TBAB
ESTIlf ATBD PBOPOB-
TIONS BASED ON SECOBD
CASE
COOKSTOWN SUBTBT
nOUBBS
1911
x*
0
0
0
1912
X**+Y*
0
0
0
1913
X-+Y<i-fZ»
X
300
286
1914
X"+Y»+Z<»
X+Y
400
390
1915
X»»+Y-+Z«
Y+Z
110
83
1916
xHy»»+z-
z
10
22
attached to each of these symbols conveniently indicates the stage of
development, as: t, infection year; d, dormant year; «, swelling stage; 6,
blister or aecial stage.
Using these symbols the accompanying tables have been constructed
showing the development of the cankers on these twigs. The first table
is based on a four-years cycle, where the swelling stage lasts but one year
before aecia are formed, while the second table is based on a five-years
cycle, where the swelling stage is continued for two years. In the third
column of both tables are entered the number of swellings which will
100 Phytopathology [Vol. 7
appe:ir oiich year, exprefwed by the h>tii1)oIs adopted. An attempt hai«
lKH»n made in the fourth eolumn to substitute values for X, }', and Z.
bailed on the proportions obtaining in the Secord case. In the Second
series the infections Ix'gun in 1914 on the growth of 1914, 1913 and 1912
were 127, 49, 0. corresponding to X, y, and Z, respectively. Adopting
JiOO as an arbitrar>' value for X, then Y Incomes approximately 100.
Z should then be zero, but since there is good reason to think that in-
fections sometimes occur on shoots of three years' standing, though none
were foimd in this case, a nominal value of 10 has \iecn assigned for Z.
Beside these* estimated pro)X)rtions in the adjoining column are
placed the actual figures of the C'ookstown sxirvey. Since these figures
indicate th(» swellings noted at one time on several successive years of
gn)wth. they may l)e used fairly to represent the swellings that would
aris<» on one y(»jir's growth in a numlK»r of successive seasons. It will Iv
MH^n at a glance tlmt while the estimated pn)portions as obtained from the
figures of tin* S<'C()rd case do not agrc^c at all with tho actual siu^ey fig-
ures in the tabic* showing a four-yt^ars cycl(». there is a strong re««em-
blance lK»tww»n thes<» simie two cohunns in tin* tabic* whore a five-yean*
cvcle is us<m1.
It is probable*. then*fore, that tlu^ tal)le giving a five-years cycle ex-
pn»ss<*s mon* nc*arly the actual dc»v«»l()pment of the disease than the table
giving a four-yc»ars cycle; in oth(*r words, the nonnal blister infection on
thc»s<* young pifH*s passi*s two yc»ars in the sw(*lling stage, and the dcvel-
opni(*nt of the <liM*ase as a whole follows this cours<*: First season, inflec-
tion; s<*cond s<*ason, donnant jK*riod; third s<»as4)n, sw(*lling stage; fourth
si'ason, swc'lling stiige; fifth and following s<*ju<ons, ac*cia.
The actuid time c*lapsing l)etw(*c*n inf(*ction and the first production of
blisti*rs acconUng to this plan of development is something short of four
vears, but sin<*e the course* of the* dis<»aM* involv<*s five* s<*asons it seems
lM*tt<*r for ])ractical n*asons to dc*signatc» it as a fiv<*-yc*tirs cycle.
The alxive conclusions as to the course* of development of the dis<»jise
on young pin«*s should Ik* fairly trustworthy inasmuch as thc*y an* Iwiseei
on a fiin^id«TaM«* numlH*r of blister cankers. On the other hand, then* i*
certain rvidftifc in Uith the (Nnikstown an<i the Se*cord cas<*s which in-
dicates that priTocious or delay«*<l devc*lopmi'nt may occur, and that
wliilt' tln' lif4'-4-vcIc* (outlined mav o!)tain in the majoritv of cas<*s, it is bv
no m<*ans to \h* n*garde<i as an invariable* nili*.
Division of Botany
Dkpartmknt of A<iiiiri i/rriiK
OrrAWA. Canada
SPECIES OF MELAMPSORA OCCURRING UPON EUPHORBIA IN
NORTH AMERICA
E. B. Mains
No species of Melampsora on Euphorbia was known to occur in the
Western Hemisphere until the present year. Collections, however, have
been made recently in the United States both upon introduced and native
species of Euphorbia. These collections, which Dr. J. C. Arthur has
kindly turned over to me for study, consist of one collection upon Eu-
phorbia eommutata Engelm. from Indiana, four upon E. robusta Small from
Colorado and Wyoming, and one upon E. Cyparissids L. from Maine.
Of these, it appears certain that the one on E. Cyparissias is introduced
and those upon E, robusta and E. commutaia are native.
Up to the present time, six Old-World species of Melampsora upon
Euphorbia have been recognized, these being M, Gelmii^res,, M. Eup-
horbicB-diUcis Otth, M. Euphorbice-Gerardianoe W. Miiller, M, HelioscopicB
(Pers.) Wint., M, Euphorbiw (Schub.) Cast, and M, Euphorbice-Engleri
P. Henn. all of which have very similar uredinia and urediniospores,
the separation being by telia and teliospores. Of these the first three
are well-marked and distinct species. Of the last three, 3f . Helioscopice
and M. Euphorbias while distinguished from the rest by well-marked char-
acters, are separated from each other, according to Miiller (1907) only
by a rather small difiference in the length of the teliospores. This dif-
ference may be a real one but is scarcely apparent in such European ex-
siccati of the two rusts as the author has at his disposal for examination.
The last, M. Eurphorbice-Engleri is a species of doubtful validity. It was
set apart by Hennings, owing to its habit of maintaining itself wholly by
the uredinia, other spore forms never having been found. This is only
known on one species of host.
A study of the North American collections of 1916 shows that they
can be readily placed in three species, two of which correspond to two
of the above and one of which appears to be imdescribed. The Maine
collection, which is upon Euphorbia Cyparissias, has teliospores which
in length and imiform thickness of wall agree very well with M. Euphor-
bice upon the same host ih Europe, while the Indiana collection upon E.
eommutata with its 31-58^* long and apically thickened (3-6/i) teliospores
agrees very well with European material of M, Euphorbioe-GerardiaruBy
• •• • •••II . ••!:••. I.. •-••
•
102 Phttopathologt [Vol. 7
the uredinioeporee of all being similar. The collections upon Euphorbia
robiuia from Colorado and Wyoming, however, are distinguished from all
other species of Melampsora upon Euphorbia by certain characters of the
uredinia and uredinioepores and are considered by the writer as belong-
ing to an undescribed species.
The North American species of Melampsora upon Euphorbia may be
keyed out as follows:
Uredinioepores 1&'23m long, paraphyses
numerous
Teliospores decidedly thickened at the
apex 13/. Euphorhia-Gerardiana,
Telioepores not or only slightly thick-
ened at the a|>ex 2 M. Euphorbia,
Urediniospores 16-29 m long, paraphyses few.. 3 3/. moniicola.
I. Melampsora EuphorbuB-Gerardiance W. Miiller, Contr. Bakt. 17*: 210.
1906.
O and I. P>''cnia and aecia \mknown.
II. Uredinia amphigenous and caulicolous, scattered, circular, 0.2-0.5
mm. in diameti'r, sul)epidermal, soon naked, pulverulent, pulvinate due
to the crowded paraph>Tje8, pale yellow, ruptured c»pidermis inconspicu-
ous; paraph>"Be8 numerous, intermixed with the spores, capitate, 16-19
by 51 58m, the wall colorless, 2-3^ thick; urediniospores globoid to ellip*
soid, 13 16 by 16-20m; wall colorless, 2-3m thick, finely and closely echinu-
late, the por(»8 ol>8cure.
III. Telia caulicolous, prolxably also amphigenous, circinating about the
uredinia, oblong, 0.2 1 mm. long, 8ul)epidennal, slightly elevated, blackish-
brown; teliospores prismatic, 9-15 by 31-60m, rounded at both ends; wall
light chestnut-brown, darker towards the apex, 1.5/i thick, 3-6m at the
apex.
On EuPHORBiAcnSiB
Euphorbia commulata Engc^lm., West side of High Lake, Noble Co.,
Indiana, June 11, 1916, II, III, C,C. Deam 2008SA, communicated by
G. N. Hoffer.
This collection, which is the first collection of a Melampsora upon
Euphorbia to be refxirted for North America, has a range in the length
of the teliospore somewhat less than that given by MQller (1907, p. 641)
for M. Eupharbict-Gerardiantt in Europe. A comparison with European
material upon E. falcata (Sydow I'red. no. 1G87) sho^-s a very close agree-
ment, however, Uith as to the ure<iinio8|x)res and teliospores. The telio-
spores are not quite so generally thickened at the apex as in the E^uropean
specimen but Mill are very (ie<'ide<lly thickened, up to 3-6^, while the
ure<iinia have the umiuiI pulvinate apiieanincc and abundant paraphyses.
Pvcnia and aecia arc not known for this 8|>ecics Init will doubtless l^e
fouml upon the same host, since M. Hdioscopue, M, Euphorbia and M
1917] Mains: Melampborab on Euphorbia 103
EupharbiaHivlcis, the three species of Melamspora upon Euphorbia whose
pycnial and aecial stages are known, are autoecious.
2. Mdampsora Euphorbice (Schub.) Cast. Observ. Myc. 2: 18. 1843.
Uredo Euphorbice-HelioscopicB Pers. p Euphorbim-exigiUB Pers. Syn.
Fung. 215. 1801.
Xyloma (Placuntium) Euphorbice Schubert in H. Ficinus Flora der
Gegend urn Dresden 2: 310. 1823.
Uromyces verrucipes Vuill. Bull. Soc. Bot France 41: 285. 1894.
Mdampsora Eup}iorbiaS''€xigiuBW. Muller, Centr. Bakt. IT*: 210. 1906
Melampsora EuphorbuB-Pepli W. Muller, Centr. Bakt. VP: 210. 1906.
Mdampsora EupJiorbuB-CyparissicB W. Muller,. Centr. Bakt. 19*: 453.
1907.
Mdampsora Cyparissias W. Mttller, Centr. Bakt. 19*: 561. 1907.
O.^ Pycnia flattened hemispherical; ostiolar filaments none.
I. Aecia foliicolous and caulicolous, circular to oblong, 0.2-0.5 mm. in
diameter on the leaves, 1-4 mm. long on the stems, orange-yellow, with-
out peridium or paraphyses; aeciospores spherical to ellipsoid, 19-24 by
21-28m; wall closely vemicose.
II. Uredinia amphigenous and caulicolous, scattered, circular or oval,
0.1-0.3 nwn. long, early naked, pulverulent, pulvinate from the mass of
paraphyses, golden-yellow fading to white, ruptured epidermis inconspic-
uous; paraphyBes intermixed with the spores, numerous, capitate, 16^20
by 31-51/*; wall colorless, 3-4/* thick, smooth; urediniospores globoid to
ellipsoid, 13-19 by 17-23/i; wall colorless, 2-3/* thick, closely and finely
echinulate, the pores obsctu'e.
III. Telia amphigenous and occasionally cauUcolous, scattered, cir-
cular to oval, small, 0.1-0.2 mm. long, covered with the epidermis, com-
pact, pulvinate, dark chocolate-brown; teliospores prismatic, 7-13 by
32-45/*; wall chestnut-brown above, lifter below, 1-1.5/* thick, not thick-
ened at the apex, smooth.
On EUPHORBIACEiE
Euphorbia Cyparissias L., Bank near Turner graveyard. Isle au Haut,
Maine. Sept. 13, 1916, II, iii, J, C. Arthur.
3. Melampsora monticola sp. nov.
O and I. Pycnia and aecia unknown.
II. Uredinia amphigenous and caulicolous, scattered or in circular
groups, circular or oblong, 0.2-2 nun. long, subepidermal, long covered
by the epidermis, pulverulent, orange-yellow, ruptured epidermis con-
spicuous; paraphyses few, intermixed with the spores, capitate, 13-21
by 32-58/*, the wall colorless, 1.5-3/* thick, smooth, the stipe solid; ured-
iniospores globoid, ellipsoid or obovoid, 13-20 by 16-29/*; wall colorless,
1.5-3/* thick, finely and closely echinulate, the pores obscure.
III. Telia amphigenous and caulicolous, circinating about the uredinia,
circular or oblong, 0.1-1 mm. long, subepidermal, slightly elevated, black-
ish-brown; teliospores prismatic, 9-16 by 27-56/*, rounded at both ends;
^ DeBcription of pycnia and aecia adapted from Dietel (1895).
104 Phytopatholoot [Vol. 7
wall liRht choatnut-brown below, darker towards the apex, 1.5-^ thick,
2-4m at the apex.
On Euphorbiace.c:
Euphorbia robusta Small, Carpenter, Wyoming, Aug. 18, 1916, II, iii»
E, T, A' E, Bartholomew 6067 {Vromyces Tranzschelii Sydow, 0, III,
also present); (Colorado Springs, Colora<lo, plains 10 miles east of citv,
Aug. 31, 1916. II, iii, E, Bartholomew, 6104 (U-pe); Palmer Lake, Colo..
Sept. 23, 1916, II, III, E, Bethel (two collections).
In the younger uredinia, es|>ccially, apparently thinner walle<l uredi-
niosjK)res are often seen mixed with the thicker. After treatment with
lactic acid, which serves to differentiate the wall from the cell contents
an<l make it stand out more clearly, the apparent difference is not to \yc
()l)serve<l.
This species is distinct from other sjx^iMes of Melampsora on Euphorlmi.
The unuhnia are abundant, large, and long covered by the epidermis
and since they (*ontain l»ut few paraphyses, after the rupture of the cover-
ing epidermis and the cscajM* of the pulverulent mass of sywres, they |x>s-
sess a Hattcnc(i ap|H*arance when contnusted with the pulvinate mass of
paraphys(\s (»f other species on Kuphorbia. The uretUnicxspores are much
more variable and larger in size than those of the other si)ecies.
No pycnia or aec'ia were found uiwn any of the collections. They are.
how(»vcr, to be lcK»ked for upon the same host earlier in the season since
the rusts of this group whose life cycl(» an* known are autoecious with all
.sjMin* forms, although Jacky (1899) working with M. Euphorbict on E
CypariMsias and Muller (HK)7 p. 449) working with the same rust on E
Peplus claim to have obtained urtnlinia by infection from teliospores, yet
Dictcl (189')) working with M. Enphorbuv on E. Cifparisitias.vLml Muller
(19t)7) with the same rust on E. exigun on the other hand have shot^ii
that pycnia and ac4'ia are prcwluce*! ujk)!! these hosts from infections with
the telio^|M»rcs. Dictcl (1S89) has also shown that M. Euphorbitr-dulci*
Srhr<H»t. has pyniia and aecia. Since th(»sc stages ar(» develo|HMl sparing-
ly, it is probable that the first workers overlcH)ked them and that M.
Euphurhyt has all s|H)r(» f<»rms. Although all the North .\merican col-
lections wcf(» <»xamincd for pycnia a.ssociate<l with the uredinia none were
found, and it is pn»bablc that all of thc.^' rusts will Ih* found to lie autoe-
cious and have all s|>*jrc fonns.
It is interesting to note that from a region in which this group has hith-
erto not U»en kiiowti so many collect i<»ns from such widely separated
are.is should have all Ihm^u made in one season. It is not so surprUing
that ^f , EupKnrhitr should Ik* f(»und u|M)n E. ('ifpnrissias.iis the latter has
lieeii brought into this country from Kuro|N* and it is likely that the rust
was intnMlue<Ml with it. It is, however, surprising that the rust
han not previously U»<»n foimd sim»<* the host is rather widely distributed.
1917] Mains: Melampsorae on Euphorbia 105
The other two species of Melampsora are upon native species of Euphor-
bia and of these M, monticola is evidently a purely American species
found upon a common western Euphorbia.* With its abimdant, large,
orange-yellow uredinia, it is remarkable that this very striking rust has
not been collected before. The other species, M. Euphorbiae-Gerardianae,
is probably native to this country' as well as to Europe, since its European
hosts, E. Gerardiana and E, falcata are not known in this coimtry. An-
other argument in favor of this assumption is that the species of Mel-
ampsora on Euphorbia have in most cases physiological races limited to
one species of host as Miiller (1906, 1907) has shown. Consequently
even if European hosts were foimd, it would be doubtful if rusts on
American species could be considered as having an European origin.
The writer wishes to express his deep appreciation to Dr. J. C. Arthur
and Prof. H. S. Jackson for the helpful suggestions and criticism re-
ceived in the preparation of this paper.
Purdue University Agricultural Experiment Station
Lafayette, Indiana
literature cited
DiBTEL, p. 1889. Ueber die Aecidium von Melampsora Euphorbiae-dulcis Otth
und Puccinia silvatica Schrot. Oesterr. hot. Zeitschr. N. 7.
1895. Ueber den Generationswechsel von Melampsora Helioscopiae und
M. vcrnalis. Forstl. nat. Zeitschr. 6: 373.
MttLLER, W. 1906 Versuche mit Uredineen auf Euphorbien und Hypericum.
Centr. Bakt. 17*: 210-211.
1907. Zur Kenntnis der Euphorbia-bewohnenden Melampsoren. Centr.
Bakt. 19*: 441-460, 543-563.
Stdow, p. and H. 1914. Monog. Ured. 3: 379.
Jacky, E. 1899. Untersuchungen iiber einige schweizerische Rostpilze. Ber.
schweiz. bot. Gesellsch. 9:49-78.
* In a commimication recently received from Mr. E. Bethel he says, **The one
[Af . tnarUicola] we have here is undoubtedly native as I found it in the high moun-
tains (above 9000 feet) at Nederland, Colo."
RECENT CULTURES OF FOREST TREE RUSTS
James R. Weir and Ernest E. Hubert
The determination of various species of rusts found on forest trees ol
the general region of Montana is a difficult task when descriptive evi-
dence and spore measurements are used. In most cases involving heter-
oecious rusts successful inoculation is considered very necessary in
determining the identity of the species imder consideration. Conse-
quently, as a beginning, an attempt was made early in March, 1916, to
secure various forms of hypertrophy formed by the bark-inhabiting Peri*
dermia. After collecting, these were placed in the laboratory in large
test-tubes with sufficient water to supply the branch or twig supporting
the infection. Many needles were always left on the branches or twigs.
In this manner the fungus in many of the infections was induced to pro-
duce Hpores prematurely and these were available for cultiu^ work at an
early date. This process also served to develop successfully the pycnial
stage of Cronartium coUasporaides (D. A H.) Arth. and Cronartium
CompUmia Arth. which stage preceded the aecial stage in both cases. In
the search for material a foliicolous rust on the needles of Larix oecidentaU$
was collected for the first time in June, 1916, at various points in Mon-
tana and Idaho. The rust was very abundant and widespread. All the
caulicolous forms of rusts on forest trees in this region and in the states
of Michigan and Minnesota as well as a number of the foliicolous forms
were tried on a variety of suspected hosts. All inoculations were isolated
at the* gnrnhoui^ at Missoula, Montana, by the use of celluloid cylinders
and (*otton plugs. The inoculated plants were sprayed daily with tap
water for a period of throe to five days following inoculation. The fol-
lowing is a summar>' of the cultures to date since the last report.^
Five pbmts of Casiilleja angusiifolia, two on May 14, 1916, and three
on May 8, 1916, were dustiMl with nei^ly develo|)ed neciospores (forced
in lnl>orator\') of Cronartium coleosporoidea (D. A 11.) Arth. (P. siatacU-
fimne ty|M») on Pinus cofUorta from Hayden I^ike, Idaho. Of the first
two plant.** one di-v(»lojMMl uredinia on May 26 and telia on May 29, the
n*n)aininf( plant dyin^ lK»fore May 24. ()f the other three plants, two
' W«*ir. J. H.. And Iltibort. K. K. SuocoAflful inorulationfl of Larix occidentalit
an<l Larix curo|N*a with Mclainp^ira l>ifcolowii. Phytopath. 6: 372-373. A^. 1016.
Weir. J. H. and HiilHTt. K. K. .\ ftucccHsful inonilation of Abien lantocarpa with
PurriniaMtruin pustulatuni. Phytopath. 6: 373. Ag. 1916.
1917] Weir and Hubert: Cultures of Rusts 107
developed uredinia on May 30, followed by abundant telia on Jime 3.
^^ •
The remaining plant wilted. Four control plants remained normal.
This cheeks the cultures of 1915.*
Three plants of CcistiUeja angustifolia were dusted with aeciospores of
Cronartium cdeosparoides (t3rpical gall form) on Pinus carUorta from
Sylvanite, Montana, Jime 23, 1916. Uredinia were not observed but on
July 17, 1916, telia appeared uniformly on all three of the trial hosts.
Three control plants remained normal. A similar result was obtained on
Castilleja with aeciospores from the gall type on Pinus contorta from Evaro,
Montana. Telia were recorded July 17, 1916, on orle of three trial-hosts.
The remaining plants died. Three control plants remained normal.
On May 23, 1916, aeciospores of Cronartium coleosporaides (blister
type) on young seedlings of Pinua panderosa from Haugan, Montana,
were dusted on three plants of CaatiU^a angustifolia. Heavy infections
on two and a light infection on one with xu*edinia on June 5 and teUa on
Jime 8 were recorded. Three control plants remained normal. This
result also checks similar cultures made in 1915.'
On Jime 21, 1916, aeciospores of Cronartium coleosporoides (typical
gall form) on IS-jrears-old trees of Pinus ponderosa from Sylvanite, Mon-
tana, were dusted on three plants of Castilleja angustifolia. The ap-
pearance of the uredinia was not recorded owing to absence from the
laboratory but on July 17, 1916, teUa were noted in abundance on two
of the inoculated plants. The other trial-host died. Three control
plants remained normal.
The results of the season together with those of 1915 demonstrate
that the various caulicolous forms of rusts occurring on Pinus contorta
and P. ponderosa in the Rocky Mountain region are the aecial stage of
Cronartium coleosporoides. It is not only possible but very probable
that the same forms on Pinus contorta and P. ponderosa as known else-
where in the western United States belong here also. An examination
of the aeciospores from galls on Pinus contorta, P. ponderosa, P. attenuaia,
P. couUeri, and P. jeffreyi, from widely separate regions has not brought
out any specific characters different from that of the material used in the
successful cultures on Castilleja. Attempts were made to infect young
leaves of Quercus rubra with aeciospores from galls on Pinus contorta and
P. ponderosa but without success.
Aeciospores Of Cronartium Comptonice Arth. on Pinus banksiana from
Cass Lake, Minnesota, were sown on trial-hosts as follows: On two plants
of CastiU^a angustifolia on May 17, 1916, with negative results; on one
' Weir, J. R., and Hubert, E. E. A serious disease in forest nurseries caused
by Peridermium filamentosum. Jour. Agr. Research 6: 781-785. Ja. 24, 1916.
> Jour. Agr. Research 6: 781-785. 24 Ja. 1916.
108 Phytopathology [Vol. 7
plant of Quercua rubra on May 18; 1916, with negative results; on one plant
of Comptonia asplenifolia on May 18, 1916, with positive results, uredinia
appearing on J\me 9 and teUa on June 17. All control plants remained
normal.
Aeciospores of Crofiartium Comptonice Arth. on Pinua banksiana from
East Tawas, Michigan, H-ere sown on trial-hosts as follows: On three
plants of Castilleja miniata on May 24, 1916, with negative resiilts; on one
plant of Qtierciui rubra on May 24, 1916, with negative results; on two
plants of Comptonia asplenifolia on May 23, 1916, and June 13, 1916, re-
spectively. Tredinirf were noted on June 13 and telia on June 23 on the
first plant and unnlinia on June 30 on the second. Two control plants
remained nonnal. Aociosfwrcs were sown on one plant of Myrica earo-
linensis and one of Myriai gale on May 26, 1916. Uredinia on Jime 13
and telia on June* 25 wctc noted on l)oth plants. Two control plants re-
niaincii nonnal.
A collection of leaves of the previous sciu^^m (1915) of Populus tremu-
loidcH lM*aring telial sori of Mclampsora rnedusw Thiini. vnis made in Pattee
Canyon two miles .southejist of Mi.ssoula, Montana, on March 12, 1916.
A p^irtion of the^sc* over-wintered leaves were placed in moist chamliers
on March 15, 1916, and a few days later upon examination of the yellowish*
brown, downy layer fonne<l on the .*<ori, it was foimd that .sporidia were
prc»>*c»nt. Thes<» fn\sh siK)ridia were ilschI to inoculate newly formed nee-
dles of lAirix (Kcid4mUiliH, Two separate inoculations were made on March
22. 1916, followcnl hy two others on March 26, 1916. Individual branches
of trees of /.. occidentalis wv.n* xh^hI for the test. On April 3, the first
inocuLition had resulted in pycnia which w(»re accompanied by small
droi>s of hyalines licjuid. On April 4 the aecia were present on the under
si<le of the ni^e<llej<. The second in(K*ulation gave pycnia and aecia on April
4. the third and fourth in(H*ulations re.nulting in pycnia and aecia on April
8. Th<» four (*ontrol plant-s of L. occidentalis as well jis the needles on the
inoculated tnM»s which wcn» not included within the cylinders remained
nonnal. -\ll the ikhmIIcs .subjected to the inoculation lx?c4une infected and
devclc»j)c<i a Lirg'j niunlx'.r of sori of l)oth .st4igi»s (O and I) of the rust.
On April 9, 1916, germinating t<ilios|K)ric material of Afelampsara
mt^iumr Thlim. on Populus tremuloidcs was sown on two small tre<*s of
hirij ruroftea. On April 26. tu\ abundant <U».velopment of pycnia were
nHMirded and on May 1 thi^ aecia ap|H^inid. The pycnia' and aecia and
their resjK»ctive sj)ori»-formH on the two sjiecies of larch were found upon
examination to U» i<lf»ntiral. The telial material failwl to infect Tsuqq
hitirophylh and Pseudotsutja taxifitlia, Tlu» control plants in all casesi
remained normal.
TeliosjMires Uuiring .s|x>ridia of MvUimpsora mctlusa Thiim. on lea\'cs of
Populus trichocarpa were coUf^tetl at Haugan, Montana, on June 5, 1916.
1917] Weir and Hubert: Cultures of Rusts 109
On June 7, these were sown on one small tree of Larix europea, two of
L. occiderUaliSf one each of Tsuga heterophylla, T, caroliniana, and Pseu-
dotsuga taxifolia. On June 15, pycnia appeared on the needles of L. eu-
ropea and on the needles of the two trees of L. occidentalism Aecia developed
on L. europea in abimdance on June 24 and on L. occidentalis on June 22.
Negative results were secured on Tsuga heterophyUa, T. carolinianay and
Psexidotsuga taxifolia. Control plants remained normal.
Teliospores bearing sporidia of Melampsora bigelomi Thiim. on Salix
cordata mackensiana from Deborgia, Montana, were sown on Larix occi-
dentalis and L. laricina on Jime 7, 1916. One small tree of each species
was used. Pycnia appeared on Jime 15 and aecia on June 18 on the
needles of L. occidentalis. On L. laricina, the pycnia were noted on June
15 and the aecia on Jime 18. A heavy infection occurred. Control
plants remained normal.
Aeciospores of Pucciniastrum pustulatum (Pers.) Diet, on Abies lasio-
carpa resulting from an inoculation with the telial stage on Epilobium
angustifolium* were sown on two plants of Epilobium angustifolium on
May 18, 1916. Uredinia developed on Jime 4, on the underside of a ma-
jority of the leaves subjected to the spores. Only the younger leaves be-
came infected. Control plants remained normal. This completes the
cycle for Pucciniastrum pustuiatum.
TeUospores of Gymnosporangium tvbulatum Kern on Juniperus scapula-
rum from Missoula, Montana, were sown on two plants of Cratcegits
douglasii on April 27, 1916. Pycnia in abundance appeared on May 8 and
11 and aecia on May 22 and 24. Control plants remained normal. This
checks a previous culture.*^
Teliospores of Gymnosporangium nelsoni Arth. on Juniperus communis
from Bonner, Montana, were sown on two plants of Amelanchier alnir
folia on May 21, 1916. On June 9, pycnia appeared in abundance, the
aecia developing on June 28. Control plants remained noVmal.
Teliospores of Gymnosporangium nelsoni Arth. on Juniperus scopulo-
rum from Bonner, Montana, were sown on two plants of Amelanchier
alnifolia on May 21, 1916. Pycnia appeared on Jime 6; aecia were
noted to be developing on June 28. Control plants remained normal
Several of the cultures are still in progress of development and as soon
as results on these are secured further information on the rusts of forest
trees in this region will be available.
Office of Investigations in Forest Pathology
Bureau of Plant Industry
Missoula, Montana
* Phytopath. 6: 373. Ag. 1916.
^ Weir, J. R. Telial stage of Gymnosporangium tubulatum on Juniperus sco-
pulonim. Phytopath. 5: 218. Ag. 1915.
A RHIZOCTONIA OF THE FIG
J. Mats
With Plate II and Three Figures in the Text *
A fungiifl possessing characteristics common to Rhisoctonia was ob-
8er\T(l to occiir in the years 1915 and 1916 on leaves, twigs and fruit of
the fig, Ficus carica, at Gainesville, Florida. Superficial, silvery to yd*
lowish white, thin mycelial strands of this fungus can be seen to radiate
loosely from yellow to dirty brown infection centers in the fig leavea and
spread over the surrounding green tissue. Later as more of the
tissue becomes discolored by the invading mycelium, the upper suifi
of these enlarged brownish areas become silvery white and dry, themyogj
ial strands become less, if at all visible, while the imder surfaoes at tlie
corresponding points remain light brown to brown and are uAially eov^
ere<t with a visible web of myceliimi (fig. 1). Small, immature wbhe,
as v»r\\ as mature bro^Ti to dark brown sclerotia attached to mycelial
threads are often foimd on the petioles and midribs of infected leaves,
but seldom are sclerotia foimd on the blades of such leaves. Loosely
woven, silky strands of the myceliimi of this fungus may be traced to
some distance on the twigs. More or less dense accumulations of sclerotia
are usually grouped on one side of the twig (plate II, fig. 2). No direct
injurious effect was observed to have been caused to the twigs by the
fimgus. The fruit of the fig may become more or less covered by the
spreading mycelium. Numerous sclerotia are then produced which
cling by mean^ of mycelial threads to the fruit and its stalk (plate II,
fig. 1)!*
Portions of diseased fig leaves, and parts of t^igs upon which mycelium
and sclen)tia were plainly visible, were killed and fixed in Camoy's fluid
and enilxHlded in paraflin. Cross-sections of the leaf tissue reveal hyphae
)H*netrating through the ^tomata into the parench>'ma (plate II, fig. 4),
while (*ross and longitudinal sections of the twig tissues do not show the
prrM'iice of a |)<.*net rating mycelium.
This fungiLH was isolated from the diseased tissues of fig leaves, where
mycelium but no sclen)tia ^i-ere present; and fnmi single sclerotia taken
from twigs. The pure <'ultures from lH)th sources were aUke in cvcr>'
n*>|MTt. I^U"e cultures of this fungus wtTt* grown easily for over eighteen
montlis on standard nutrient agar, (*orn nu^al agar, and sterilised green
1917] Matz: Rhizoctonia of Fiq 113
Of the previously described specieB of BMzootoiiia the follcwing should
be considered here:
Dr. Peltier* in a recent publication on parasitic Rhizoctonias in Amer-
ica summarizes as follows: "At the present time there are recognized in
America two species of truly parasitic Rhizoctonias; the common form
RhieocUmia Solani Kilhn {Corticum vagum B.& C.) , widely distributed and
occurring on a great number of hosts; and R. crocorum (Pers.) DC,
with a limited distribution on alfalfa and potato tubers. A third Rhizoc-
tonia, Corticum ockralettcum (Noack) Burt, is found on the leaves of
pomaceous fruit trees, while a fourth species isolated from damped-off
^
Fia. 3. Myceuuu of Rhieoctonia HiCBOBCLEaOTiA
A, Short chains and a group of short cells from the margin of young sclerotia.
X 333.3. B, Vegetative mycelium from a young agar culture on the left, and from
ma old culture on the right. X 333.3.
onion seedlings is of questionable parasitism." In discuseing the growth
characters of a number of strains of Rhizoctonia, Peltier* makes the fol-
lowing statement (p. 370) regarding the species isolated from damped-off
onion seedlings: "The strain from onion produced sclerotia which were
entirely different from those of other strains in that they were small
(0.5 to 1 millimeter in diameter), perfectly round, bright colored, and
developed submerged in the medium." None of these latter characters
have been observed in the Rhizoctonia from the fig. The sclerotia of
D America. Illinois Agr, Exp. Sta.
114 Phytopathology (Vol. 7
this fim|i^«< are roundish or oblong, from 0.2 to 0.5 millimeters in diameter,
an» not bright colored and arc produced freely on the surface of but not
in the medium in culture.
The effects and manner of attack of Corticium ochraleucum (Noack)
Burt on pomaceous fruit«, as described by Stevens and Hall,^ are similar
to those of the fig Rhizoctonia on its host. However, the sclerotia of the
two fungi are entirely different and the |)erfect stage of the former has
not so far been found in connection with the fig Rhizoctonia on fig trees.
I>uggar* in his recent paper gives sufficient evidence that the conmion
Rhizoctonia in America is Rhizoctonia Solani Kiihn. This Rhizoctonia
has Cftrticum vagum B. & C. as its perfect stage. In comparing the
Rhizoctonia of the fig with the Rhizoctonia Solani Kiihn, obtained fmni
l)ean .s<»edlings here, the two show pronounced differences on storiliied
l)ean plug and agar slant cultures. On each medium the sclerotia of the
first are white at first, turning dark brown with age, and remain small
and more or less glolK)se; the sclerotia of the second are white at first,
turning light brown to brown or dark brown and are ver>' irregular in
size and fonn (plate II, fig. 6). Mycelium and sclerotia from pure
niltunw of the fig Rhizoctonia and from R. Solani Kiihn were plac*od on
moist and growing fig leaves and twigs. Both fimgi killetl areas in the
leaf tissue but R. Solani did not produce any scleortia on the infected
parts. Two separate flats of cowpea seedlings w<»n» inoculated with the
two fungi. R. Solani killed 90 per cent of the scredlings, while the fig
fungiL'^ did not produce any injur>' to the young plants.
Shaw* in his account of a Rhizoctonia which he found on jute, nml-
lK»rry, cotton, and cow])ea, and which he apparently misnames Rhitoc*
tenia Solani Kiihn, illu.strates a fungiLs which is similar to the Rhizoc-
tonia of the fig in sevenil res|x»cts, i.e., the nonnal occurn»nce of itji
niuiiennis, com|>!iratively small, rounded wlerotia on the tips and stems of
it*< n»s|KH'tive host plants. However, the diameter of the sclerotia of
ShawV Rhizoctonia is alnmt half of that of the fig Rhizoctonia; the
color <»f the .*(<!l(Totia of the former is black, while that of the latter is
bn»wn to dark bn>wn. Shaw's illustrations (plat(» IX) indicate a distinct
I'ort^^x in tin* srlcrotia of his Rhiz^x'tonia. which fonnation is al>sent in
* St«'vrnJ«. F. L. ami II all. .1. (I. HypocnoM* (if poinarcHMiK fniit, North Caroltna
Agr. Kxp .^tft. Hi-p 190e<»:.T.>. Tft Ki. fig?* II IV VMl.
* DuKKar. H. M. Uliizoctonia rroroniiii (|Vr*4.i I). (\ and It. Holaai Kuha
iCortiriiiiii vaKurn H. tV (' < uith iiotrM on othor mimm-Icm. Ann. MisHouri Hot. (lani.
3: 4o:i-4,> vm:»
•.*<haw. K. .1 K MMq»hMloKy an«i para«»iti**in of Uliizoctonia. Mc»m. Dt'pt. Afpr.
Iniiia 4: Ml. I'.MJ.
The (mmiii" Uhizortonia in lii«li:i Mi*in. I)i>pt.. Akt.. Iiulia.7: no. 4. I9IS.
1917] Matz: Rhizoctonia op Fig 115
those of the fig. The mode of origin, as described by Shaw, of the sclero-
tia of the first, has not been observed in the fig fungus.
In culture the sclerotia of the fig fungus develop from dense masses of
short hyphae and thefe the young sclerotia are usually s\u*rounded by
short chains and groups of ovoid, short, sometimes elbowed cells.
Zimmerman^ in describing Corticum javanicum on Coffea arabica,
Coffea liberica and several other plants, mentions the occurrence of a
sterile mycelium and small (0.15 to 0.3 millimeter in diameter) white
sclerotia-like bodies (weisse Kugelen) in association with this Corticum.
He states: "dieselben treten sowohl auf der Ober — ^als auf der Unterseite
der Zweige auf imd sind ausserdem auch namentlich haufig an den Friich-
ten zu finden;" but no mention is made of its occurrence on leaves. In
describing the effects of this fungus on the host, Zimmerman says, *'Die
unter den Kugelen gelegenen Pflanzenteile sterben — zmn mindsten in den
nahe der oberflache gelegenen schichten — ^ab und erhalten eine dimkel-
braune bis schwarze Farbung;" but no direct injiu*ious effect from the
sclemtia of the fig fungus was observed on the branches of its host. Re-
garding the "Kugelen" of Zimmerman's fungus he states: "vertrocknen
sie einfach an den zweigen, auf denen sie sitzen." No mention is made
here of the change in color which is a dark brown in the sclerotia of the
fig fungus at maturity.
Edgerton* described a limb blight of the fig, due to Corticum Icetum
Kars. This disease is characterized according to Edgerton's illustrations
nnd descriptions, mainly by the conspicuous fruiting layer of the fungus
associated with the diseased parts of the host, but no mention is made of
any Rhizoctonia occiu-ring on fig trees afflicted with limb blight.
Kuijper® describes a leaf disease of Coffea arabica and Coffea liberica
under the name of **Zilverdraadziekte der Koffie," in Surinam. There
is a striking similarity in the character, the manner of attack, and effects
of the sterile fungus which causes the Silverdraad disease on Coffea, to
the Rhizoctonia of leaf blight of the fig. In describing the Coffea fungus,
Kuijper does not mention nor illustrate anything which approaches a
semblance of the dark brown sclerotia which are commonly foimd in
connection with the fig fungus on its host and in pure culture. The
sclerotia of the latter are not identical with Kuijper's "hyphenkluwens."
These forms occiu* also in his cultures. He states: '*0p plaatsen, waar
veel, zijtakken ontstaan strengelen deze zich door elkaar, zoodat op de
^ Zimmerman, A. Ueber einige an Tropischen Kulturpfianzen beobachtete
Pilze I. Centbl. Bact. Abt. II, 7: 102. 1901.
•Edgerton, C. W. Louisiana Agr. Exp. Sta. Bui. 126: 13, pi. VII, fig. 1. 1911.
• Kuijper, J. De Zilverdraad-ziekte der Koffie in Suriname. Dpt. van den
Landbouw, Suriname. Bui. 28: — 1912.
116 Phytopathology (Vol. 7
Waderen l)e8rhrevon khiwens ontstaan." In sonic of his cnlturos Kuijprr
obtained **h>'phon-opconhoopinf2:en zoo st<»rk, dat bijna lH)lfomiigi'
lichaampjos van 14 2 ni.m. doorsnedo ontstaan, die l)est{ianuit oon l>otrpk-
kclijk los lu^phenvlechtwcrk.'* Apparently these Imdies never Uvonie
in Knijper's cultures a.s compact and colored as do the sclen)tia of the fig
fimicus on various media and on the host.
Brooks and Sharpies'' in their work with Corticmm salmonicolor B. &
Br. (C jafanicum Zini) descrilx* four forms in which the fim^^s appt^ars
on ruhl)er trees as follows: **a pink incrustation on the branches or main
stem; .... white or pale pink ])ustules arranged more or l<»ss in
lines parallel with the branches; .... part of the* fim^s on the
exterior consists of white or pale pink stnmds of a cobwel)-like texture,
which run irrei^ilarly downwards over the surface, the strands l)einK
.»H)metimes .<o <lelicate as to be overlooked: .... finallv th«»re is
the Necator stag(* consists of oninf?e-n*d (not pink) pu.*^-
tules alK)Ut «)ne-(»ij?hth inch in diameter.'* In their pure cultures of (\
salmonicolur, the alM)ve authors observed clamp coimections in old a>
well as young cultmes, a pink to a bright ros(» coloration and, **aggn'ga-
tions of hy])hae .... n^st^nbU'd a numlH»r of clos<»ly attached
Necator ])ustul<»s." Practi<*ally iiorn* of thes<' characters were ol>served
in the fig fungus.
From the precciling di.*<cu.'<sion it follows then that the Khizoct(»nia
of th(j fig leaf blight is different fnnn all the true s]H»cies of Hhizo<*toni.*t
previously descrilxMl and thus it should Ih» considcnMl as a new organism
which can briefly Ik» descrilHMl as follows:
Rhizoctonia microsclerotia n. sp.
Srh-rotia su]MTlicial. small 0.2 to 0..') millimeters in diameter, white
when yoimg. brown to dark brown at maturity, nearly homogenous in
sinK'lure an<l color, sub-glolM>s<', free from t\ift4'd myi*elium, not ."^nHMith
UMmlly single, sometimes conglomerated.
Vegetative hy])hae 0 to H^i wide, first hyaline and granular, bniwn an«i
more f»r le**-* empty with m.aturity, septate.
Ilab. On living leaves, branches antl fruit of tin* cultivated fig, Ficus
cnrioi, ( lainesvilk*, Florida, V. S. .\.
»• Hrookii, F. r. an«l Shan)!**". A. Pink •liHoa.-MV Dopt. of .\Kri. FcvleratM
Malav St!it«'H Hul -M : 1 -27. fig 1*». 101 1.
1917] Matz: Rhizoctonia op Fig 117
Rhizoctoii^ia microsclerotia Sp. Nov.
Tuberculis super ficialibuSy minusculiSf 0,2 to 0.5 millimeters in diameter,
primum aUns deinde fusco-bruneis (intus idem quod extus), similibus
fere forma et colore intus ac extuSj sub^lohosis, floods myceliaribus defidenti-
bus, non gUxbris, saepius singulatis, non-nunquam conglomeratia.
Hyphis 6-8 li, latis, primum hyalinis granulosis deinde bruneosis,
septatis.
Hob. in foliis, ramis ac fructibus vivis Fid caricae cuUae, Gainesville,
Florida, Am, Bar,
University of Florida Experiment Station
Gainesville, Florida
Pl.ATK II. KlIlZtM'TONIA MirKOHC'LKROTiA ON FlO
Vui. 1. H. iiiitTusrhTotiii rovcrinK ii tuiic ami fruit of the fif(. X 1.5.
FiK- •• FifC twiK defoliHt(*<i l>y the fuiiKUM. XumorouN M*lerotia are arcumii-
\iitvi\ on ono Hide of the twif^. X !..'>. «
Ki<i. 3. Tlir(M'-wiM'kHH>l(l rultun*^ on Mtrrilizetl hotui {mmIm. U. mirrosclerotia od
left. H. H^ilani Kiihn on right.
Fi(i. 4. CroH.H MOf'tion of (liHo;i.'4o<l fifc hMif showinx prnotration of myootium throuich
Htomata at a and h.
h'ui. 5. Sf»rtion throuich a Arlcrotiuni of K. mirrosolorotia.
Via. Ti. Srl«*rotia from a Hk hranrh. X 2r>.
11*^
1917] Sherbakoff: Buckeye Rot of Tomato 121
any fungal growth. When the affected fruit is kept in an enclosure with
high -humidity, especially when the fruit approaches maturity, the fungus
which causes the rot may be observed commonly on its surface (fig. 2A).
The rot occurs on the fruit in all stages of its development, beginning
almost invariably at the point where the fruit touches the ground. Nat-
urally, the fruit touches the ground most commonly with the blossom-end,
on which account this rot often appears as a peculiar form of blossom-end
rot and for which it sometimes has been mistaken.
NAME
The disease is known among some of the growers on the East Coast as
water logged fruit. This name cannot be adopted for the rot because it
is misleading and because it has no other advantage in its use. The name
brown rot though it would often describe the disease, ought not be used in
reference to the disease, first, because there are even more frequent cases
when the color of the rot is not distinctly brown, and second, because the
name- has been applied already by Bancroft to an apparently different
rot of tomato fruit.
Some of the tomato buyers use the name buckeye, in reference to a rot
of tomato fruit. The writer was unsuccessful in finding with certainty
what particular rot is called by that name, but his indirect information
and observations indicate that the name refers to the rot under consider-
ation. This name describes very well the most striking feature of the
lecaon on fruit affected with the disease, namely, its broad zonation, in which
case the lesion indeed much suggests the eye of a large animal. The name
also has not been used in literature before in reference to any disease of
a similar natm-e. Therefore, it is suggested that the common name, buck-
eye, should be used in reference to the tomato fruit rot described here.
OCCURRENCE
The rot was foxmd by the writer for the first time in January, 1915, at
Goulds, Florida. Soon after the first observation and during the following
three months it was found in every tomato field of that locality on the
prairies — ^low marl lands usually under water during rainy summer months.
The whole district is known as Redlands and lies at the extreme south
end of the Florida East Coast, namely, south of Miami.
The same rot was observed by the writer in 1916 on the West Coast
also. In April of that year it was found in a field near Bradentown, and
in May in a shipment of tomato fruit received by the writer from a field
near Palmetto. In both instances the tomatoes were grown on the com-
mon low hammock land of that vicinity.
122 Phytopathology (Vol. 7
The writer also found amon^ some old specimens of tomato fruit pre-
><erved in fonnaliii and kept in the laborator>' at Gainesville, two ^fieci-
niens which by external symptoms and microscopic examination pnived
to Im^ affected with this same rot. One of the specimens had a lal)el indi-
cating that it came from Little River, near Miami, in 1911. The other
8|)ecimen l)ore no label.
All these observations show that the rot occurs in South Florida on
both coasts where it is common in low fields even during a comparatively
dr>' season, such as that of 1916. No actiail observations were made of
its occurrence in other parts of the state, but judging from its common
presence in so widely separated parts as the east and west coasts and on
soils s<j different in character, one might safely assume that it is much more
generally distributcnl than observations indicate.
HISTORY
The ])rescTvr(i s|>(»ciin<»n of the rot found in the lal)orator>' and previ-
<)iL*<ly mentioned, shows that the (lis<*as(» was pres<Mit in the state at least a^
early as 1911. An incpiiry among tomato growers of the East Coast aL«o
indicates that it hjis existed ther(^ a long time.
Pathological literature, with one excerption, contains no reference to any
dis<»as(» of tomato fniit similar to the rot \mder consideration here. The
organism which caus<»s this rot is clos(»ly related to I^hytophthora infe^ftarut
but the latter fung\is is not the same as the one which causes the rot herein
de.**crilxjd and the disease* prochiced by I\ infcstans is also dilTf»rent lie-
cause it affects all aerial parts of th(^ plant and the fungus always pnNlu(^*>
abuiulant conidia on the surface of the afT(Tted parts of the fruit.
The previously mentioned exception is a short note by Bancroft* on llio
Brown Rot of the Tomato, in which is descrilKMl briefly the s>'mptoms.
orcurreiice, economic im|N)rtance, and the method of transmission of the
dis<»a.*«4*. No illustrations of any kin<l at'company the article to assist in
identifi<*ation of X\\v dis4*a.*<<» and in general it is t<K) brief to judge with
rerlainty whether tlu* tomato fruit rot n'|H)rted from Knglan<l is the same
as. or difTen*nt from the rot found in Florida. It apjX'ars tlmt the rot of
tomato fniit re]>orted by Banrroft has thc»s4' two ft^itures similar to the
burkeye rot : « 1 ) The dis<»as<* occurs «)nly on the fruit ; (2) the fungus asscK
ciateil with tin* rot in Kngl.ind is closiHy related io the fimgus which
cttUS4»s the r«»t in Florida.-
* Hanrrnft, C. K. The Immn nit ni toninto. Jour. M. .\in'. (London) 1$: 1012.
1910
» Whether It i* ri'.illy thr simw fiiiiKU«t or ii«»t it is* iiiipoMiMc to determine, be-
rnuM* ulioitt III** fuiiKa?^. Daiicroft iiicrrly nay^ "the tiHMiieit of the endosperm and
emhryo" nf thr afTrrte*! need "contain fungal hyphae, which from their cbarmclert
1917] Sherbakoff: Buckeye Rot of Tomato 123
The points in which the two rots appear to differ are as follows: (1) In
describing the rot in England, Bancroft says: "A fruit which is infected
first shows discolored patches on its surface; these usually run together,
so that the whole or almost the whole surface of the fruit becomes discol-
ored." The rot observed in Florida as a rule appears in form of one,
rarely more, gradually spreading, often distinctly zonate spot, usually at the
blossom end of the fruit; (2) Bancroft from all his observations on the rot
says this in regard to the way in which the disease affects the fruit: "This
infected seed .... is known among the growers to be capable of
germinating and is reported to produce plants which always bear infected
fruits. These facts, coupled with the results of microscopic examination
of the seed, suggest that the hyphae may be capable of existing in the seed
in a resting condition, becoming active when the seed germinates, and keep-
ing pace with the growth of the plant until the fruit is formed." That
is, Bancroft evidently records no other method of infection of the fruit
than the one suggested in the above citation, while it is evident that in
the case of the rot in Florida the infection starts from outside the fruit.
From the above citation it is also evident that the rot reported by Ban-
croft is associated rather with the use of infected seed (the plants from in-
fected seed "always bear infected fruits") while in our case it evidently
is associated with the infested soil, and only the fruits which touch or
nearly touch the ground are affected with the rot.
On the whole, it seems that the disease described by Bancroft is differ-
ent from the rot of tomato fruit found in Florida, and the latter should
therefore be considered as a new one to the literature.
ECONOMIC IMPORTANCE
The data at hand will not justify any precise statement concerning the
extent of damage caused by the rot. But considwing that the tomato
crop is the most important of all vegetable crops in Florida, and consider-
ing also, that the rot was actually observed to affect up to fifteen per cent
of the fruit in the field and up to ten per cent of the fruit in-transit, it is
evident that the disease is important.
CAUSE
Repeated isolations from the tissues of tomato fruit affected with the rot
invariably yielded one and the same fungus, usually in pure cultures di-
appear to be hyphae of Phytophihora omnivora.^* Hyphae of a number of fungi
belonging to the same family do appear under similar conditions alike, the fungus
causing the rot in Florida included; and yet they are quite different, but to differ-
entiate them much more than appearance of the hyphae is needed.
124 Phytopathology [Vol. 7
rectly fn«n plantings. The firbt isolations were made from the material
collectoil on the Kast Coast, in Janiiar>% 1915; the last in May, 1916, from
material obtained fnnn the West Coast.
The isolations wvre made hy planting small bits of the affe<*ted tissin*:*
(from under the epidennis and after the fruit was disinferted in menMirir
ehloride sohition, 1 : KNM), for aUnit fifteen minutes) into either {X'tri
dishes with a suitable mediinn (<*orn-nieal and oat agars wen* most com-
numly used for this pur]K»se) or into test-tulH»s with sterilized lioan |>o<l<
or oat agar. Dilutions of swarm-si>ores wen' <Mnploved to make certain
that the cultures were pure. This was an «»asy proci»dure l)ecauw the fun-
gus readily s]><»rulat(»s and the swarin-s|>ores an* produced from mature
conidia ^swannsporangia) in a very short time and practically under all
conditions of th<* lalNuatory (under favorable <u)nditions s^'ann^pores
were (>roduced in some instances eight minutes after the culture was plaoed
in frrsh water). \ very successful proc(Mhire here is t<» place a ffood-
sized pitM'c of )M'an-|)o<l (*ulture (two or nion* weeks ol<|) of the fuilglM
into a sterilize* I watch-glass, with a frw <'ubic rciitimet«Ts <if Hterilind
water in it. for al>out fortVH'ight hours; then, after the culture in thtt
watrh-glass is washed a few timrs in fresh striilizrd wat<T, it is left in the
watrh-glas?* with a frw cubir crntinu'trrs of wat«T for half an hour. In
the first two days the lM*aii-p<»d cultun* usually produces a great masB of
ronidia; in half an hour or an hour after it is rinsed, these im'annspcrBngMl
will liU*rati' int(» tht> water a ronsidcpablr number of swarm-spores; this
water with numerou'^ >warni-**|)ores in it '\< to be us«m| then for poured*
]>late dilutions made in the ordinary way. ( *orn-meal agar was com-
monlv :ind >uiTes««ful!v umm| I»v the writer for these dilutions.
• • •
Inortilution** of tomato fruit. detarh<>d from the f)laiit, and not detached,
anil of all •'tani's from very yoinm to n'd.i'i(M' with ])ure cultures of the
fungu**. inv.'triably n*>ulted in re])rodurtion of the rot (fig. 2). The inocu*
lalion-i wen* made by plaeing sm.all bit< of the culture and water Hiu^pen-
^•ion*' of *warm-^pores upnn wounded and un wounded "Surfaces of the fruit.
In the (';i*«e of '*w:inii-^]MiM- iiiiH u]:ii ii»ii a pjerr of th«' fungus culture wa.s
dri»]»[H-d intii a l.iii:«' j.ii nearly full nf -lerilizid w.ati-r and then the fruit
wa*i pla<-<'d mtii thr watii. Deiarlu'd liiMiato fruit-* float in water and un-
iletaj'heij fruit ran Im- eniiVi-nii'iil Iv ]jla«-ed in '•ueh :i way that only a part
of It vill bi- in till- water.
huM'ulatinns iiitii wmmdeij funr ne.trly :ih\;iy- -liowed the rot in tWt»nty-
four hour-. Iiiorulatmn- iiiti» unwounrjrd fiuif. wliethiT myrclimn or
-Warm— |H>rf- wiir rni]»l«»\i li. •^nmrinnr*. ohn\M-d the rot in lwi»nty-four
hour- .ifier thr iri«»iu!:iMf»fi \\a- maiji". but oftrn the inferiion c<»uld not
III- ijfTi itril until thn-r t»i •vi-n four ♦lay- later. \\ ]»ii*s**nt no explana-
tii»n 1- f»lT'ri'.| for t*'is variatM»n mi the ineubalion |M'riod.
126 Phytopathology (Vol. 7
Th<» fungiis has Ikvii j^rown and studiiMl on various niotlia parallel with
other organisms related to it, sueh as Pythium debaryanum He««*., Py-
thiacystis ciirophihcrn Sin. &, Sm., and s<»veral s|)eoies of Phytophthora,
P. cnctorum (I>»lKTt & Cohn) Sehr. and P, er y throne ptica Pethybr. in-
rhided. Tliis eoniparative study shows plainly that the fungus causing
till* huekeye rot of tomato fruit is different from tlM» others. Its peculiar
tufte<l gn»wth on the corn-meal agai*^ in plates, is one* of the differentiating
characters, <»s|KM*ially valuable* for an easy s<»paration of this fungus fnmi
the others (fig. 3).
Kxamination of the literature also indicates that this fungus luis no
U»en ])reviously di'scrilMMJ. However, evidently th<' same organism wa
isolated iH'fore. though from another host; but it has Ikvii considere<i to
Im^ the siune as the fungus of the lemon brown rot, namely, Pythiacysth
nirnphthorn Sm. A: Sm. The ref<»renee hen* is made to the organism iso-
lated by II. S. Kawcett^ an<l by 11. K. St< vens from the bark of citnis
ln»es in Flori<la affected with the foot rot.
The writer's C4)mparativ<» study of the fungus of the buckeye n»t of
tomato, of the fungus of the lemon brown rot. P. citrophihoro,iind o{ the
fungus from \\\v citrous f(M>t rot in Florida shows that the tomato n>t
organism ami the one of the citrous foot rot are morphologically aiul cultur-
ally identical and that it is <listinctly different f nun the true /^ ri7ro/>/i//ifwci.
.V fungus was recently isolatcvl by the writer from a lupine stem n>t
which IS evidently the sam<' as the one of the buckeye rot of tomato fniit.
liHM'ulations of tomato, sw<H't |M'p|M'r. watermelon fruit, of lemons and
of iuIhts of Irish |>otat<H's with pure cultures of the s<'veral s|K»cies of
Phvtophthora, ])reviou>lv mentioned, of the Pvthium, IMhiacvsti** and
thn*^' strains of the fungus of the buck(\ve rot, inclu<ling one strain isolated
by H. K. Stevens from <lis<»as4M| bark of a citrus tre<' in Florida affected
with the f(H)t rot, sh<»w that certain organisms distinctly diffen»nt in their
mor()hology may affect the same host and pnxluce more or lc*ss similar
effects. The fungus of the Inickeyt* rot prcMhu'cd a rot of all the parts of
the plants hen* n'ftTnMl to.
These inoculations thus indicate that the organisms which do attack the
Sjiiiie plant or ]>lants cannot on this basis alone )><* considen*d identical
with each <»ther.
• Thi* t<»rri-fiH*al »ic»r i?* inudr l>y hratiii)! .'lO teraiiiM of r<>rii-int>nl in inOU rr. of Ui*-
ttllfil wnt4'r at fio ('. for an hour, then filtrriiiK tli«' li(|ui(l through i\ filter paper,
adWiiiK to It \'» itrninn of agar and the rfciuiml amount of <iiHtill(Hl wat(*r to KrinK
vohinu' of thr h<|tii<l to KWM) rr.. ('(Hiking the Muh?«t:uire in a doiihlr lK>ilcr until ml!
agar in (li!*S4»lv('4l. ((Mihng <ioun to .V) i\. .vhhng the white of an egg. nutoelaviii|t
at l.'* |KMin(l» of preHxure for alnMit fifteen iiiinute»<, filtering through a filter pa|M*r.
tiihing and auto* laving a* Ix^fore
• Fa^eetl. II. S. Tho known (lixtriluition of I*vthiae\>ti?« citniphthora an«l it»
pfohahlc relati(»n to mal di goninia of rit^l^. IMiytopath. 6: tWK 07. 1915.
1917] Shebbakoff: Buckeye Rot op Tomato 127
On the basis of the data obtained in the course of the writer's compara-
tive study of the fungus of the rot of tomato fruit and of the other fimgi
related to it, it is considered an undescribed species which may be briefly
characterized as follows:
Phytophthora terrestria n. sp.
Myceliiun at first continuous then septate; conidia usually terminal,
sometimes intercalary, mostly oval, papillate at apex but variable, 42.5
X 30.5 (36-46 X 24-35)|i| germinating mostly by swarm-spores; swarm-
spores asymmetric, with two cilia on one side, 9.5 (&-ll)|i, when in resting,
globoid stage; chlamydospores common, mostly globose, 34 (SO-40) /jl; oogo-
nia common in old cultures on steamed bean pods, globose, 22 (19-24)|i
with the stalk evidently penetrating through large, nearly globose antheri-
dium (fig. 4); oospores globose, 20 (18-21)^; colonies on corn-meal agar, in
petri dish, peculiarly tufted.
Hab. Parasitic in tomato fruit causing buckeye rot, in bark of trunks
of citrus trees causing foot rot, in stems of a Lupine causing stem rot, and
apparently in low soils; in Florida.
Phytophthora terrestria sp. nov.
Mycelio priuo continuo deinde septato; conidiis fere terminaltbus aliquando
ifUercalaribiia, pterumque ovoideis apice papiUatis sed vcUde variabUibua,
4£'5 X S0.5 {36-46 X 24-35)yL, per zoosporos fere germinatis; zooaporis
aeymmetris, laieraliter S-ciliatis, 9.6 {9-ll)yL diam. quum quieti, globosi statu;
chlamydosporia vtUgarilms, plerumque gkbostbtAa, 34 (30-40) fi diam.; ooganiis
globosis, 22 {19-24) ii diam., radicibua per amplum sub-globosum antheridium
aperte penetrantibus; aosporis globosibus 20 {18-21)ii diam.; cohmiis in agaro
Zeae Maydis farina, in petri patera, euo genere criakUi.
Hab. paraaitice in frudibue Lycopercici esculenti efficiens "bitckeye rot,**
in tmcis Citri efficiens ''foot rot," in caulibiLS Lupini sp. efficiens **stem rot,"
et aperte in humiUbus solis, Floridensibus, Am. bar.
CONTROL
No direct control experiments with the buckeye rot have been con-
ducted, but the fact that the rot occiu's almost invariably only when the
fruit touches the ground, or is very close to it, naturally suggests staking
the tomato plants to prevent it. The staking would hold the fruit high
enough above the ground to keep it from attack by the fimgus.
The fact that the fungus attacks the fruit from the groimd, coupled
with the fact that the rot, once it starts, progresses under normal condi-
tions fairly rapidly (often nearly the entire fruit may become affected in
Phytopathologt {Vol. 7
1917] Sherbakopf: Buckeye Rot op Tomato 129
three days after inoculation, the rate depending evidently on the tempera-
ture, moisture content, and maturity of the fruit), also suggests a method
of control of the rot while in transit. Here it probably would be very ad-
vantageous to keep the fruit for a few days after it is picked before pack-
ing it for shipment. All fruits that were infected in the field would de-
velop the rot sufficiently to be detected by the packers and thus be thrown
out without contaminating the rest of the fruit, which then could be
safely packed and shipped. But the practicability of these methods has
not been tested.
SUMMARY
1. The buckeye rot of. tomato fruit is common in certain places on the
low lands of the east and west coasts of Florida.
2. It occurs only on the fruit that touches or nearly touches the groimd.
3. It is caused by the fungus Phytophthora terrestria n. sp., which is also
found on other hosts.
4 It causes considerable injury to the fruit in the field and in transit.
5. Staking of the plants in the field and keeping over the picked fruit
a few days before it is packed seem to be practical methods of its control.
UNrvERsiTY OP FLORroA Agricultural Experiment Station
Gainesville, Florka
Explanation op Fig. 5
Af conidiophores from culture on hard, oat agar, eight days old. X 100. B,
conidiophores and C, intercalary conidia; from the same medium and of the same age
as A, D, conidiophores from the fimgous growth on an artificially inoculated (up-
per) and on naturally (lower) affected and nearly mature tomato fruit. Ej oospores
within the oogonia with the basal antheridia evidently penetrated by oogonial
stalk. Ff germinated chlamydospores. G, t3rpical and H, various abnormal forms
of conidia, produced on surface of a mature tomato fruit affected with the buckeye
rot. I, resting and germinating swarm-spores. Ky motile swarm-spores (fixed in
Flemming's fluid and stained with eosin). L, conidium germinating by a single
germ tube. M, two conidia germinating by many germ tubes (evidently each
swarm -spore which did not escape gives its own germ tube). B-L X 500.
NOTEWORTHY PORTO RICAN PLANT DISEASES
F. L . Stevens
The following diseases are of interest to pathologists of the United
States, either because they are new, or because they are little known
diseases of important crops belonging within the territory of the United
States or are caused by fimgi of special interest for one reason or another.
CoFFEB. Pellicvlaria koleroga. This fungus occurs in great abun-
dance, especially in the lower altitudes in Porto Rico. In destructiveness
and general appearance it gives an impression somewhat like that of pear
blight with many branches killed and the dead leaves matted and hang-
ing by their fungous attachments. The fungus forms thin skin-like
membranes over the lower leaf surfaces and has thread-like growths
leading down the petiole and adjacent branches. In habit it strongly
resembles Hypochnua ochrohucua Noack as it occiu*s on apples in the
Carolina mountains, and the writer long has had the suspicion that the
two fungi are related.
Hemileia vdstatrix. This destructive fungus, though not seen in Porto
Rico, is worthy of mention on that account. There is a report that it
was imported into the island, almost immediately recognized, and due
to the vigilance of th^ experiment station officials was so thoroughly
eradicated that not a specimen has since been collected. The following
quotation from a letter from Mr. May, in charge of the Porto Rican
Experiment Station, is worthy of record.
"When the station was first established in Porto Rico we were carry-
ing on some experiments in the Carmelita cofifee plantation, five hours by
horseback above Ponce, imder charge of J. W. Van LeenhoiBF. In 19(^
or 1903, I do not remember which, it was before I came to the station,
Van Leenhoff got some little coffee trees from a Dutch warship that
brought them from Java. After he planted them out he noticed that
they had what appeared to be Hemileia vcistatrix. Mr. Van Leenhoff
had been a coffee planter in Java and was pretty sure of the fungus from
such investigation as he could make. The matter was commimicated to
Washington and L. A. Clinton, of Connecticut, was sent down to inves-
tigate the trouble. In the meantime Van Leenhoff took every precau-
tion, destro3dng all the plants and all material that might have in any
way been connected with them. Clinton spent some weeks at the Car-
1917J STBVBNS: PORTO RICAN PLANT DISEASES 131
melita but oould find no traces of the fungus and as none has appeared
since, Van Leenho£f doubtless made a 'dean up.' Since that time great
care has been exercised with all coffees brought from foreign ooimtries."
StUbeUa flavida. The characteristic circular leaf spot caused by this
fungus is common in the higher altitudes, but is never fo\md in low re-
gions. The fungus is of imusual scientific interest on account of its
problematic relationships. It is by no means limited to coffee but is
found on numerous hosts.
Sugar Cane. Lepiosphceria Sacchari, This fimgus is quite commonly
present as a destructive leaf spot.
Palm. Graphida Phoenicis. This fimgus, which is of uncertain re-
lationship and is common as a minor pest in northern greenhouses, is
found in the open on several species of palms in Porto Rico. It has been
noted particularly upon the date palm and the hat palm. The hat pahn
is a very profitable plant, a single leaf being worth about ten cents. Fre-
quently trees arc seen with all the leaves closely covered with Graphiola
and lines of diseased tissue reaching long distances through the petioles.
The financial damage is done to the young, as yet imfolded leaves, the
fungus penetrating them and rendering them worthless.
Melicia furcaia. This is present sometimes to such extent as to largely
cover the leaves with its black coating. It has not been observed on cul-
tivated palms but occurs in great abundance on wild Thrinax ponceana
on plants of marketable size. As yet, however, there seems to have
bepji no attempt to make a commercial enterprise of shipping the bcau-
tifiil young palms which spring up spontaneously in such profusion in
Porto Rico.
Auer$waldia palmicola. This was noted only a few times, but in those
instances affected nearly evcr>' leaf and leaf-segment on the tree.
Bread Fruit. Oredo Artocarpi, The inmiense leaves of the bread-
fruit harbor numennis fungi. The crop, if it can l)c called such, is not
of high value, and this r\tst, though interesting, is not of much economic
signific4ince.
Corn. PhyUachora graminis. Mention liaj< Iwen ina<le of this in
Phytopathology by Miss Nora Dalby. The disease was \%nde-spread in
Porto Rico and must Irnve l)een to considtTuble exti»nt injurious.
Bean. I>imerium grammodes. This striking fungus which occupies
and covers the veins of the affected jmrts of the leaf with i\^ conspicuous
black perithecia is common on several grnera of h^gtuncs, among them
cultivate<i beans.
GuAVA. Meliola Pauiii. The Mark s|>ots of this fungiu< an» almost
universally pn»sent where\Tr the host is foimtl. Then* is but liitli*. if
any injiuy.
132 PHYTOPATHOLOGY [VOL. 7
Aschersonia, What is taken to be one of this genus sometimes occurs
profusely on the lower surfaces of guava leaves, giving them a livid scar-
let color, conspicuous to a considerable distance. The fimgus appears
to be growing upon a scale insect, probably Alerodes citri.
Cephaleurus tnrescens. This algal parasite is of special interest. The
spots show well from both below and above the leaf, the leaf tissue being
killed. This alga, also widely known as the cause of a serious tea dis-
ease, is present parasitically on a large numl)er of Porto Rican plants.
GoNDULE. Uromyces dolicholi. This universally present leguminous
plant is almost always rusted to a slight degree, sometimes badly rusted.
Mango. Meliola Mangiferce, Meliola is very common and wide-
spread but apparently not injurious. Gloeosporium mangiferct is often
abundant and injurioiu^, especially upon the fruits of the finer varieties
of the mango. The effect in general is much like that of the bitter rot
on apple.
Sweet Potato. Coleosporium Ipomcece, Rust is common on this and
other Ipomoeas, but iLsually not t^) serious extent. In one field at Pres-
ton's Ranche near Nagualxi, infection was general and serious. Each
sorus was also parasitized by a Fusarium-like fimgus, rendering it white.
Grape. Physopella vitis. This is a rust of cultivated grape. Three
collections were made in Porto Rico but all from the same vines, namely,
at Patillo Springs. The rust was pn»,sent in quantity suflScient to make
it injurious. The writer knows of only a few grape vines in Porto Rico.
If there were more \'\nv» perhaps there would have been more collec-
tions of this rust.
Peantt. Vramyces arachidis. Only one collection of this rust was
made. Indec^d, the peanut plant is not very common in Porto Rico.
Canxa. Pxicciniii Canruz. The rust on Canna is common on both
wide and cultivated Cannas in all parts of Porto Rico; on nearly evcr\'
one of the host plants. Sometimes the rust is so abundant as to appear
fairly destructive.
I. NO A Lai'rina. Microstroma sp. This host, an important leguminous
coffee shade tn»e, is frec|uently nmrh infej^ted ^^ith an undescribed Micro-
stroma which ca\L**es large witches br(M>ms.
Pitiiecolobium Saman. MicroHtrama sp. This le^(uminous tree is
being intro<hic<Hi into Porto Rico through the (efforts of the experiment
station, and in the sihmI Ih»(L< and propagating lyoiU it is frequently heavily
infest4Ml with an ap|wrc»Jitly un<le-»<cril>ed sp<H*ies of Microstroma.
Pasi»alum. MyriftgctutsfHtra .**]). Th(» Pa.»*pahmis of yard and pa*-
tUH' <ift4*n lH»ar this vcr>- intc»n»sting fungiu*. Infertion is usually gen-
eral throughout thr plant, that is, if one part is infected e^ich leaf is likely
to Ik* invad(*d.
1917] STEVENS: PORTO RICAN PLANT DISEASES 133
Melia. Psendoperonospora sp. Two collections were made, in
widely separated parts of the island, of this fungus. The damage is
probably slight. The four fungi last mentioned will be described fully
by Dr. Lamkey.
Ficus. Kuehneola Fid, This rust was very conspicuous and pres-
ent on a large number of species of this genus. Certain large trees at
the proper period of the year were repeatedly covered with the rust and
often small shoots a foot or so high would have each leaf completely
rusted.
Cotton. Kriehneola Gossypii. Very little cotton is raised in Porto
Rico. This rust was foimd in considerable abundance in one field, though
probably not doing much damage.
Manihot. Uromyces Janiphce. Only one collection of this rust was
made in Porto Rico, though the host is very common there under culti-
vation, and the rust has been looked for repeatedly. .
Sooty Molds. The sooty molds familiar to Northern pathologists
on the orange and CamelUa and to lesser degree on many greenhouse
plants, aboimd in Porto Rico. There seems to be no speciaUzation to
hosts and they grow indiscriminately upon all hosts. If it be a large
plant as a mango tree that is primarily infected, the fungus, spread
doubtless by rain, is found growing upon every kind of plant beneath
the tree. Very Uttle evil eflFect is noticeable other than the imsightly
condition produced. The statement is frequently seen that these sooty
molds exist upon insects or insect secretions. Such organic matter cer-
tainly favors them and increases their luxuriance, but the sooty molds
are not entirely dependent upon insects and insect products and may
aboimd without them.
While many of the Porto Rican sooty molds are much like, perhaps
quite like, the sooty mold of the orange, other sooty molds diverge more
or less from this in character, yet show similarity enough among them-
selves to allow them to be classed in the same group.
The writer has so far refrained from using latin names for these fungi.
The orange and camellia sooty molds are best known under the generic
name Meliola, and the generic conception of Meliola was broad enough to
receive them imtil about 1892. Gaillard's monograph of MeUola then
showed that these forms clearly differ greatly from the typical Meliolas
in essential details, and he excluded from Meliola the tropical sooty
molds. Today any student of Meliola would, I think, agree with him.
Meliola with its capitate and mucronate hyphopodia, its characteristic
mjrcelium, perithecia, asci and spores constitutes a well-defined genus
from which the sooty molds with their bead-like mycelimn, peculiarly
shaped pycnidia and perithecia diflFer widely. The fact that they should
134 PHYTOPATHOLOGT [VOL. 7
be excluded from Meliola is clear. Jiist where they should be placed is,
however, not so clear. They comprise a rather well-defined group which
in Porto Rico certainly consists of a large number of species on many
hosts. This group perhaps coincides in limits with what Sacoardo in his
earlier volumes calls the sub-family Capnodiese and which he in his
foxuleenth voltune calls a tribe. This tribe is not recognised by Lindau
in the Naturlichen Pflanzenfamilien/but is included in the Perisporiacea.
Clements gives the Capnodiacese family rank. Amaud in 1911 in his
monograph '^Contribution a Tetude des Fumagines" Ann. Ek^. Nat.
Agr. Montpellier, places them for convenience in the ''Spheriaoees dic-
tyosporecs." Many writers today place the better-known of these forms
of the habit of the old Meliola Citri, Meliola CameUcB, and so forth,
in the genus Capnodium. Thus we have C. HeUatumf C. Mangtferum, C.
Coffe<Bf C. brasiliense, C. footie. Others place these forms in the genera
Pleosphseria, Antcnnaria, Apiosporiimi, and so on. It is not the pres-
ent purpose of the writer to attempt to determine the generic limits
here or the status of these species, but rather to call attention to the
rich mass of material occurring in the tropics, which may well be called
the Sooty Molds and in the main belong to the Capnodies of Sacoardo.
Illinois U.niversity
Urbana, Illinois
PYCNIAL STAGES OF IMPORTANT FOREST TREE RUSTS
James R. Weir and Ernest E. Hubert
With Two Fiqubes in the Text
The discovery on September 29, 1916, at Darby, and on October 8,
/916, at Bonner, Montana, of abundant exudations of pycnospores on
swellings of Pinus panderosa (fig. 1) and Pinus contorta caused by Cronar-
Hum Comandrce Pk. somewhat alters the impression that the pycnia of the
caulicolous species of forest tree rusts appear only during the spring or
early summer months. This \musual appearance of pycnospores seems
to be of sufficient importance in the life history of this rust to be re-
ported at this time together with further facts concerning the pycnial
stages of Cronartium coleosporoides (D. & H.) Arthur,^ Cronartium Comp-
tonicBy^ Arthur and Cronartium cerebrum (Pk.) H. & L. Spaulding* in his ac-
coimts of the white pine blister rust {Cronartium ribicola Fisher) states that
the pycnial stage may be found early in the spring or at almost any season
in late summer or fall, he having foimd them on the hosts in November
within a month after placing in the greenhouse. The collections of
Cronartium Comandroe on Pinus ponderosa and P. contorta bearing the
pycnial stage were of the spindle-shaped tjrpe of swellings and bore in the
central portion cankerous corrugations of the current season's (May,
1916) aecial eruptions. The pycnial drops appeared on the freshly
swollen areas at either end of the spindle-shaped hjrpertrophies bordering
the ruptured areas. This conforms to observations made on a collection
of the pycnial drops made near Bonner, Montana, on May 22, 1916,
from lesions of Cronartium ComandrcR on Pinus ponderosa. The pycnial
exudations, consisting of a clear, sticky, sweet liquid with a large num-
ber of minutely pyriform spores in suspension, appear as large or small
drops issuing from minute blister-Uke swellings in the epidermis of the
infected tissues. Measurements of the pycnospores of the various col-
lections made agree closely with those made by Boyce,* (50) 3 to 4 m by
• Weir, J. R. and Hubert, E. E. Recent cultures of forest tree rusts. Phyto-
path. 7: 106-109. 1917.
• Spaulding, Perley. The blister rust of white pine. U. S. Dept. Agr. Bur.
Plant Ind. Bui. 206: 27-28. 1911.
. The white pine blister rust. U. S. Dept. Agr. Farmer's Bui. 742: 12.
1906.
• Boyce, J. S. Pycnia of Cronartium pyriforme. Phytopath. 6:446-447. D.
1916.
136 Phytopathology [Vol, 7
3 to 7 M (3 by 4). They are characteristically pyriform, of a pale turtle
gre<'n color^ issuing from minute openings in the epidermis and are pro-
duced from 8ul)epidermal pycnial stromata of irregular outline. Boyoe
states that the pycnospores are hyaline. An examination of both i«io-
lated and massed spores indicates that they are colored, thoiigh faintly.
The pycnial stages of Cronariium coleosporaides and Cranariium Comp-
lonicB have, up to the present, remained unknown. Mention of the dis-
cover}' of the stage has been made in a previous report* but details were
not given at tliat time. In the period from April 4 to 15, 1916, abimdant
pycnial exudations containing pycnospores were obtained from galls of
Crofiartium coleosporaides (fig. 2) on Pinus ponderosa and P. cantoria.
These pycnial dmps were found in the field and were also forced in the
lalK>ratory at a much earlier date than produced in nature.* Out of a
total of 32 galls, 28 proiluced pycnia by the forcing process, several of
these* lat^r producing aecia. The pycnia of Cronartium coleosporaides on
galls develop similarly to those of Cronariium Comandras with one marked
diff(»n»nce, that is, their appearance on old galls and legions. The pycno-
spores of Cronariium Camandrce apparently develop but onoe on the same
tissue prc'ceding the appearance of the aeciospores. The production of
aecia kills the infected tissues which are included in the aecial ruptures.
The tissues lK)rdering this area are invade<l by the mycelium of the
fungus, produce swellings, and give rise to pycnospores, either in early
spring or in late simimer and fall, whenever sufficient time has elapsed
from the last prcxluction of aecia. In the cases recorded the pycnospores
appt^ired in the same season following the pnxluction of aecia, with only
five months intervening, but not from the identical area from which the
pycnia were pr<Hluced. In Cronariium coleosporaides the pycnospores
an* pr(Kluce<l on ol<l galls previously ruptured as well as on unruptured
infert<»<l tis^nies. A description of the pycnia of Cronariium coleosporoideM
on galls follows:
Pyrnial iitroma in irroi^ularly Hhapini aroAi«, more or less scattered or anasto-
moflinK. raulirolouM, subepitlemial, forming minute, blister-like swelliDgs when
mature <m unruptured infecte<i tiMuofl and iiwuinK from cracks in the bark of old
le^ionfl; exuding a dear, ffueet. stieky fluid in which the pycnosporee are suspended
forming dropfi of a ca<imium yellow to orange color when first appearing, l»ecomins
clear aM the Hpore maj«ti KettleM to the lower end of drop, and orange to brick-red
ufxm drying. PyrnoxporeH hyaline, mostly spherical, occasionally ellipsoid or
olM»vttte i,V) 1.5 to U.O M by 1.5 to 3.7 /i (2.5 by 2.5).
*i<idg<'v%a>. HnlMTt. Color ^t:indlird•« and color nomenclature. PI. 32. 1912.
• Phyt<.path 7: Un't 10l». 11U7.
' The -aine.
1917] Weir and Hubert: Pycnial Stages of Rusts 139
from the field bore remnants of the pycnial exudations and at the same
time were producing aecia in abimdance. The pycnial drops when fresh
have a slightly darker yel'ow color than those of Cronartium coleosparoidea,
and like the latter appear in the bark crevices of old galls as well as in
minute blister-like swellings on the yoimger unruptured tissues of in-
fected areas. When old and dried the pycnial exudations are difficult of
detection on the surface of the infected tissues and have a brick-red color.
From observations made on Cronartium coleosporoides on galls both
in the laboratory forcing experiments and in the field, it is determined
that the aecia follow the pycnia in the same season, usualy from eight to
sixteen days apart but not appearing upon the identical areas. The
galls shown in figure 1 were collected at Coeur d'Alene April 1, 1916,
and were placed in test-tubes with water on April 3. From April 4 to 9
abundant pycnial drops were produced. From April 11 to 14 the galls
began to show evidence of aecial eruptions and from April 16 to 17 aecia
appeared. With a longer time-interval between the two stages this is
what has been observed to occur in the field. Spaulding^ has observed
in the case of Cronartium ribicola that the pycnia precede the aecia by a
short period.
A very interesting fungus frequently foimd during the past season in
connection with the pycnial exudations and also foimd accompanying
the aecial stage of the caulicolous rust is a species of Tuberculina which
may be referred to T. maxima Rostrup.* This fungus was found at-
tacking the pycnial and aecial stages of Cronartium Comandrce occurring
on PiniAS ponderosa and P. contorta, the pycnial and aecial stages of Cro-
nartium coleoaporoidea on galls of PiniLS ponderosa and P. contorta, and the
pycnial and aecial stages of Cronartium cerebrum on Pinua bankeiana.
This species of Tuberculina attacks the stromatal layer as well as the
fruiting bodies and erupts in powdery, Hlac to nigrosin masses through the
epidermis of the host tree. This fiuigus is considered by Tubeuf • to be of
some economic importance although Lechmere^® does not concur in this
conclusion. The writers have not found it occurring in sufficient abun-
dance to place any importance upon its economic possibilities.
Office of Investigations in Forest Pathology
Bureau of Plant Industry
Missoula, Montana
» U. 8. Dept. Agr., Bur. Plant Ind. Bui. 206: 27-28. 1911.
• Tubeuf, C. von. Ueber Tuberculina maxima, einen Parasiten des Weymouth-
skiefem-Blasenrostes. Biol. Abt. f. Land- u. Forstwirthschaft. 2: 169. 1901.
* Tubeuf, C. von. Recent observations on the blister rust of Weymouth pine.
Naturw. Ztochr. Forst.-u. Landw. 12: 484-491. 1914.
^* Leohmere, E. Tuberculina maxima, a parasite on the blister rust fungus of
the Weymouth pine. Naturw. Ztschr. Forst.- u. Landw. 12: 491-498. 1914.
REVIEWS
Edible and Poisonous Mushrooms. By W. A. MurriU, Afisistant Director
of the New York Botanical Garden. Handbook, IGmo., pp. 71,
large colore^l chart, figs. 47. Published by the Garden. Price $2.
This work is primarily of interest to users of mushrooms. Tlie chart
contains very good illustrations of 30 species of edible mushrooms, and
of 17 species of poisonous ones. A few of these are of interest to the
forest pathologist, Ix^cause they frequently attack the wood of living trees.
Geo. G. Hedgcock
PHYTOPATHOLOGIC'AL NOTES
Sew hosts for Razoumofskya americana and R. ocddenlalis abieiina,
Rdzoumofskya americana Nutt, has l)een previously reported on PinuM
coniorta, P. banksiana, P. jwnderosa, and P. jeffreyi. On April 12, 1915,
a specimen was received from J. K. Haefner of the Sifiki>'OU National
Forest, Oregon, on Pinus attenuata. On Septeml)er 5, 1916, the writer
collected l>oth staminate and pistillate plants on Pinus aUenuaia in the
Oregon Mountains, Siskiyou Forest. This indicates that the species may
1h» exi)ecte<l to occur on any of the yellow pines. (See writer's note.
Phytopath. 6: 414. 1916.)
Razoumofskya occi4ientalij< nhietina (Kngelm.) (V)ville, the large form on
Abies. ha.»< Ihh^ii rei>orted on Abi4's concoloTy A, grandis^ and A, magnificat
During a trip on the Oater National Forest, Oregon, in Septeml)er, 1916,
the writer <*ollertcHl iM>th staminate and pi.stillate plants on Abies nobilis
and .4. amabilis.
James K. Weir
Lighttiing injury to kale. In a previou.s article (Phytopath. S: IM.
191')) the writer, jointly with (SillH^rt, reiK)rt<»<l ol)8erv'ation8 upon light-
iiiiit; killing of |N)tat<H\«< and <*ottoii. As sliown, the plants in the stricken
zone die j)roniptly over a somewhat eireular area usually fnmi one to
three nnU in diiiineter. Further evidence* of this sort of injur}* in Wis-
* Ilo«lKt*'><4c. (>. (f. N(»t("* on florne dineimiM of trees in our national foreau. V.
Phytopath. 6: I7«». I'M.V
Wi«T, J. K. .MiMtlctfM* injury to conifi*rH in the northwest. U. S. Dept. Afr.
Bui. :WM): 33. 1910.
142 Phytopathology [Vol. 7
of the storm were rather small and all had the appearance of having beeo
struck by some strong force and flattened to the ground, and the three
plants which seem to be inside the circle and have survived were ahnost
completely covered with earth. The rest of the plants in the circle had
a powder-burned appearance and all shrivelled up and disappeared within
a week or ten days." The area as shown in the figure was roui^y
circular, about twenty to twenty-five feet in diameter. This accords
closely in all respects with the effect4s of the lightning strokes as we have
observed them in Wisconsin potato fields.
It is to be hoped that others may record, as they have opportunity,
evidence of such injuries imtil we have a fuller imderstanding of these
matters. From the e\adence at hand, similar injuries may be expected
with sugar and garden beets, and possibly carrots, with the varioiis mem-
bers of the cabbage-kale groups of vegetables, turnips, radiahee, and so
forth, and probably with the allies of the potato, such as tomato and egg
plants. Special attention may also well be given to possible li^^tning
injury of legumes in view of Sitensky's observations. (See Abstract,
SSeitschr. Pflkr. 8: 148, 1898), which have come to my attention since
our former article. He reports a lighting stroke in a Bohemian alfalfa
field when the plants were in blossom. The next day the plants were
wilted down in a circular area about 5 meters across.
It is noteworthy that no case is recorded of like injury with any of the
Gramineae, although it would seem that, since great areas are occupied
by the grains and grasses in the northern states, lightning must often
strike in such fields. When more evidence is at hand conoeming the
varying liability of such plants to injury in nature it will pave the way
to some very interesting experimental work to determine the reasons
why such variations occur. These may conceivably be associated with
differences in the character or habit of aerial parts, with the character or
distribution of the root systems, or with the relative electrical conductivity
of the different plant tissues.
L. R. Jones
Puccinia glumarum. In May, 1915, the occurrence of Pucdnia glu-^
marum (Schmidt) Rriks. and Henn. was reported for the first time in the
Tnited States. It is so well known in Kurope and certain Asiatic countries
thiit it 8 discover^' at ho many widely distant points in the western states
led to considerable speculation as to time and method of introduction into
America. Although we are Htill lacking definite information on these
points, it is now definitely kn<mii that an examination of herbarium
s|XM*imen8 at the New York liotanic (iunlens indicates the fact that P.
glumarum was collected in this country as long ago as June, 1892, when
1917] Phytopathological Notes 14S
C. V. Piper reported it as P. rubigo-vera on Elymus americanus and a
month later as P. rvbigo-vera on Bromus hookerianusy and distributed it
under numbers 41 and 206. These specimens were found at Seattle and
Everett, Washington, respectively.
Other American collections of Puccinia glumarunij made prior to 1915^
were by E. T. and E. Bartholomew in August, 1913, at Billings, Mont.,
on Hordeum jubatum under number 4369; E. Bartholomew on Hcrdeum
jubatum at Rock River, Wyo., in August, 1911, under nmnbers 1063
and 3763, and by A. O. Garrett in 1907 and 1909 in Utah and distributed
as P. rubig(M)era imder numbers 138, 191, and 192.
It would thus appear that Puccinia glumarum has been present in
America at least twenty-five years and possibly longer.
H. B. Humphrey
Newton B, Pierce, The death is annoimced on October 13, 1916, at
the age of sixty years, of Mr. Newton B. Pierce, formerly pathologist in
the Bureau of Plant Industry.
Mr. Pierce in his early manhood was engaged in the lumber business,
in partnership with his brother at Ludington, Mich. He was from a boy
interested in natural history and spent a great deal of his time in the
woods. At first he turned his attention to economic entomology, and
very early developed into an excellent entomological artist. He was
sp)ecially gifted in the field of draughtsmanship,* particulary in the
delineation of plants and insects. Desiring to advance his knowledge
along entomological lines he went to Harvard and took up some special
work in entomology. Conditions were not favorable for the best work,
and he relinquished his entomological studies, and, after casting about,
decided to take up work in plant pathology under Dr. Volney Spaulding,
of the University of Michigan. Largely through Dr. Spaulding's influ-
ence, Mr. Pierce became intensely interested in plant pathological studies.
He developed into a keen observer and a thorough-going investigator.
In these early days there was comparatively little pathological work
carried on in this country, and those engaged in it were few in number.
The pathological work of the government had only just been inaugurated.
About 1887 or 1888 there appeared in CaUfomia a serious grape disease,
which spread rapidly and caused immense damage to the vine industry
of that state. Early in 1889 the disease had become so virulent that the
Section of Vegetable Pathology in the U. S. Department of Agriculture
decided to undertake an investigation of the trouble. In casting about
for someone to take up this work Professor Spaulding of Michigan was
communicated with. He reconunended Mr. Pierce, and Mr. Pierce was
app>ointed. Mr. Pierce proceeded at once to Santa Ana, CaUf., and made
144 Phytopathology (Vol. 7
that place his headquarters. He began at once a careful field study of
the disease. After six or eight months of field studies he decided that
he wanted to go abroad in order to get a line on the diseases of the grape
in the south of France and in Italy. He went abroad at his own expense,
and was away six or eight months. Upon his return he renewed his
investigation of the grape diseases, and eight or nine months later pub*
lished his valuable report on the California vine disease. Mr. Pierce
continued grape investigations and took up other lines of work.
Gradually the laboratory at Santa Ana grew into one of the most
important branches of the plant pathological work of the government.
The name was changed to the Pacific Coast Laboratory, and Mr. Pierce
was put in charge. He remained in charge of this work until December
31, 1906, when he resigned.
During Mr. Pierce's work on the Coast he conducted important in\'e0ti-
gations in the California vine disease, leaf curl of the peach, walnut bli|^t,
and the diseases of the grape.
Mr. Pierce was a man of quiet and unassuming habits. He was pre*
eminently an investigator, and preferred to work alone. A careful study
of the record of his accomplishments on the Pacific Coast will show that
he was an indefatigable and earnest student.
B. T. Galloway
Pathological greenkoxue. On the basis of the presentation in Pryto*
PATHOLOGY for February, 1916, of the need for controlled greenhouse
conditions in Imtanical research (Potter, Alden A. The control of experi-
mental conditions in ph>'topathological research, p. 81) funds have been
secured for carrying out these ideas in connection with the investigations
of the Bureau of Plant Industr\' on the cereal rusts, and it is hoped to
have at Ic^st one unit of this apparatus in operation in Washington within
the present year.
Cereal rtui nurvey. An extensive barberry and cereal rust field survey
is projecte<l for the coming spring and summer by the Office of Cereal
Investigations, Bureau of Plant Industry.
PernonaU, Dr. J. L. Wcimer, formerly assistant in the Department of
Botany of Purdue l*niversity, I^fayette, Indiana, has been appointed
scientific assistant in the Bureau of Plant Industr>', effective February 14,
to take up work on the diseases of sweet potatoes and other truck crops.
Mr. Moses Irvine, assistant in the Department of Botany and Plant
Patholog\\ Tniversity of Minnesota, has recently Ixren appointed assistant
in Plant Patholog>' in the Kansas Agricultural College, Manhattan, Kan.
REPORT OF THE EIGHTH ANNUAL MEETING OF THE
AMERICAN PHYTOPATHOLOGICAL SOCIETY
The eighth annual meeting of the Society was held in Barnard College, Columbia
University, New York City, N. Y., December 27-30, 1916, in conjunction with the
American Association for the Advancement of Science.
About ninety members were present and a program of eighty-three papers was
presented, the abstracts of which appeared in the last number of Phttopathologt.
Twenty-four new members were elected, making a total of three himdred and
sixty-seven.
Joint sessions were held with Section G of the American Association for the
Advancement of Science and also with the Botanical Society of America.
The following officers were elected:
Presidentf Dr. Mel. T. Cook, Agricultural Experiment Station, New Bnmswick,
N.J.
Vice-PrenderUf Dr. Charles Brooks, U. S. Department of Agriculture, Wash-
ington, D. C.
Councilor for three years t Prof. H. S. Jackson, Purdue University Agricultural
Experiment Station, Lafayette, Ind.
One of the Chief Editors of Phytopathology for three years, Dr. G. P. Clinton,
Agricultural Experiment Station, New Haven, Conn.
AssociiUe Editors, J. B. Rorer, Port-of-Spain, Trinidad; Prof. H. P. Barss, Ore-
gon Agricultural College, Corvallis, Ore. ; Dr. Geo. M. Reed, University of Missouri,
Columbia, Mo.; and Dr. H. A. Edson, U. S. Department of Agriculture, Wash-
ington, D. C. *
The Society decided to hold its next annual meeting at Pittsburgh, Pa., in con-
junction with the American Association for the Advancement of Science, Deoeniber
28, 1917, to January 2, 1918.
AMENDMENT TO THE CONSTITUTION
Article II, Section 1, line 1, after the word "include" the word sustaining was
inserted.
Article III, Section 3, line 1, after the word "become" a sustainxhg life member
by paying one hundred dollars in ten conseciUive annual payments was substituted for
"a life member upon the payment of fifty dollars."
BEPORT8 OF COMMITTEES
The Committee on Common Names, consisting of F. C. Stewart, G. P. Clinton,
F. L. Stevens, E. C. Stakman, and W. A. Orton, presented the following report and
recommendations:
Carrying out the instructions of the Society at the Columbus meeting, the Com-
mittee sent copies of the partial list of conmion names which it had prepared to
about 200 memoers of the oocietv for their criticisms. In consequence of the sug-
gestions received the Committee has made some changes in the list and 30 names have
been stricken from the list. The revised list contains the names of 17 host plants
(alfalfa to cabbage) and 92 diseases. The adoption of this list is recommended.
The Committee reconmiends further that this work on common names be con-
tinued under the following plan:
146 Phytopathology [Vol. 7
•
(1) The American Phytopathological Society shall officially adopt a list of
common names of plant diseases.
(2) There shall t>e a oermanent standing committee of the Society, called the
Committee on Common Names of Plant Diseases, consisting of five membera, one
memlier being elected by the Council each year to serve five years. The Cammittee
shall elect its own officers. The meml)crs of the present Committee (1916) shall
retire in the order of the length of service of each. A vacancy on this Committee
shall be filled temporarily by appointment of the President of the Society, the ap>
pointee to serve until the next annual meeting, at which time the Council shall
elect a meml)er to complete the unfinished term.
(3) This Committee shall prepare and present to the Society for official action
at the regular annual meeting a list of common names of plant diseases, and at each
succeeding annual meeting a supplementary list of names may bo presented. At
least six weeks l)efore the annual meeting the Committee shall submit a preliminary
list of names to all nieml)er8 of the Society for suggestions and criticiam in writing.
At the l)eginning of the annual meeting the revised list shall be conapicuousiy
posted to invite further HUggentions and criticisms. The list as finally revised by
the Commit tei» shall In* presented to the Society at the same annual meeting for
final adoption. The list of names officially adopted at each annual meeting shall
l¥» printed in Phytopathouhjy in the n^port of the meeting of the Society. The
official list may lx» amended undrr the same procedun*.
I'lKin motion the report was arcepte<l an<l the recommendations adopted. Later,
in connection with a diHcu.HMion of the propo.sed list of common names which the
Committee ha<i prepan»<l, a motion to reron.Hider the action in regard to the report
of the (^ommittfH* wa^^ adopted, and a moti<m waM made and carried that the (*ora-
mittet* be in»<tnirte<l to submit to the entire membership of the Society by mail the
list of common nafncM which it approve<l and to re<]ueHt a vote for or against each
name propose<l, a three-fourths majority of the total membership being necessary
for adoption of any name. \ motion was also adopted authorising the Committee
on (\»mmon NamoM to change the above plans to make them accord with the
alNive motion.
('. \^. Kdgerton was ap|M>intcMl tm the C<»mniittee in place of O. P. Clinton, whoee
term expiriMl. Since the close of the meeting F. C. Stewart and F. L. Stevens have
resigne<l and the PreHident has ap|M)inted in their places (i. H. L>'man and J. B. 8.
Norton.
The (*tfmmittt'e ttn Wnt^ and Means, consisting of L. H. Jones, C. L. Shear, C. W.
Kdgerton, H. S. Jackson, and J. T. Barrett, di<l not report, the Chairman being
al>f«ent.
The Committee **ri liiUliography, consisting of L. U. Jones. C L. Shear, and R. A.
Harper, mafli'no fonnal re^mrt. C. L. Shear stated that the (Committee had been
unable, after two years' effort, to wcure the fifty subscribers necessary for publica*
tion of the propos4*<i card in<lex to phytopathological literature. It had been the
exfMTtation of the Conunitte<* that a large profHirtion of the agricultural colleges and
expf*rinM*nt stations would subsrribe for the index, but only ten such sulMcriptioofl
hail lH*4*n obtaintsl. The informal report was accepted and the Committee
disrhargfHl.
Tht Cnmmittee nn the Schirrinit: (%tUection of Fungi, consisting of C. L. Shear,
3. ('. Arthur, and \. (i. Johnson, made a report of pn»gresfl to the effect that the
(*urat<ir of the Herbarium had offfred to take whatever steps were feasible to ae-
romplir«h th«* pur|M>se desirc^l by the S<N<iety. It is hop<Kl to |>erfect the details and
havr th»" matter satisfactorily atten<le«l to during the prosent year.
Thr Cirmmittte on InMitutional Stamiardisation, c<msisting of H. S. Reed, H. H.
Whi'txd. an<l H H. Fulton, premMittMl no re|>ort and re<pi(*st<Ml that it be discharfed.
Thr r»'<|U»*st was approve<l by the Society.
1917] New York Meeting 147
The Committee on Summer Meetings, consisting of Paul Murphy, E. C. Stakman,
and Donald Reddick, presented no report, the Chairman being absent.
The Committee on Pure Culture Supply Laboratory, consisting of C. L. Shear, L.
R. Jones, and G. P. Clinton, made a report of progress, and stated that an item had
been introduced in the appropriation bill for the Bureau of Plant Industry for the
next fiscal year to cover the inauguration of this work. Whatever is accomplished
in this direction during the year will depend upon the fate of this appropriation.
tbeasurer's report
Receipts:
Balance from 1915 $655.64
Dues of —
268 regular members for 1916 $804.00
72 sustaining life members 720 .00
9 members for 1915 27.00
1 sustaining life member 100.00
Overpayment 3 members 9.00 1,660.00
Exchange from 8 members. 1 .22
Interest 15.64
Excess transfer from Society funds 5.00 $2,337.50
Expenditures:
Approp'riation to Phytopathology 1915, and 1916 400.00
Clerical work (inc. $14.80 for 1915) 73.43
Printing abstracts, stationery, etc 185.07
Secretary's traveling expenses to attend Columbus
meeting ^ 44.28
Mimeograph work for F. C. Stewart 15 .00
Supplies 7.75
Exchange on checks .40
Excess dues from 3 members 9 .00
Rebate for No. 1, Vol. V, exhausted .50
Sinking fund for Phytopathology 501 .58
Telegrams 1 .55
Transfer to Phytopathology acct. members 969.00 2,207.56
Balance $129.94
FINANCIAL STATEMENT OF BUSINESS MANAGER OF PHYTOPATHOLOGY
Receipts:
Balance from 1915 $47.91
Advertising guarantee 1915 150 .00
Subscriptions and sales Phytopathology 620.38
Annual dues transferred account —
9 members for 1915 $ 18.00
268 regular members, 1916 536.00
72 sustaining members (1 excess) 360.00
1 sustaining member in full 50.00
1 member who paid $10 5.00 969.00
148 Phytopathology |Vol. 7
Sales Phytopatholoot direct 18. 10
Neiberg subscription (sent to W. A W.) 3.25
Extra illustrations in PHYTOPATHOLoaY, O'Gara and
Hotaon 11 .28
Annual appropriation from American Phytopathological
Society for 1916 200.00
Interest on deposits .84
Interest on mortgage, 6 months 15.00 $2,035 75
Expenditures:
Cooke portrait S26.00
Separates from Phytopathglooy 41 .05
Stationery, express, postage (Dr. Reddick) 35.00
Phoenix files .57
Illustration of horse-chestnut 10.00
Readjustment of dues one member paid to publishers . . 3.00
Readjustment Neiberg subscription, paid to treasurer. . 3.25
Insurance on stored stock 5.40
Manufacture of Phytopathology:
No. 6, Vol. V, balance from 1915 $65.80
No. 1, Vol. VI 432 82
No. 2, Vol. VI 348.52
No. 3, Vol. VI 338.78
No. 4, Vol. VI 260 92
No. 5, Vol. VI 165 82
No. 6, Vol. VI, $147.30, pending 000 00 1.612.66
Williams A Wilkinn mii«cellaneou8 bills for postage, etc. 78.98
riericalwork 70.43
Reimbursement American Phytopathological Society
account over-transfer 5.00 1,892 24
Balance $143.52
These accounts were referred to an auditing committee, consisting of A. D.
Selby. R. Kent Beattie. and K. C. Stakman. The Committee reported that they
had examine<l the arrounts and found them correct, and the reports were adopted.
RKHOLl'TIO.VH ADOPTED
A Committee, conHixting of II. .\. KdHon. .\. I). Selby, and John L. Sheldon.
was ap|H>intfil by the Society to draft resolutionH in reganl to the deaths of two
members, \V. \. Mnrtin an<l Vungyen Young, and alno of Professor T. J. Burrill.
The following rem»hition» were pre.«M»nt«»<l:
"HtfiUvfd, That whereaM in th<* death of W. .\. .Martin, of lloulton. Maine, and
VuriKyfii Young, of Shanghai. China, the American Phytopathological Society has
btMMi <b'prive<l of t>»o of itn menib«*rH. the Society reconU itM sincere regret at the
taking of theMc gentlemen.
"That whcrea-* in the death of Profe?*Mor T. J. Burrill. of the Cniversity of Illi-
nois, uho (ir<«t (ictnoii**trat<Hi the exintenci* of bacterial di.'^eaMes of plants, there has
bci-n rrninv***! .m eminent lea<ler in botanical and phytopathological research, the
Anteriran Phytopathological Society i'XpreK?«e?* itM nincere regret at the departure
of tliM eminent Kcholar and teacher, anci records itM appreciation of the service
renderwi our science by his researches.
1917] New York Meeting 149
"That these resolutions be filed with the records of the Society and printed in
Phytopathology."
The following resolution was also passed by the Society:
"Resolvedj That the Society express its deep appreciation and gratitude to the
local Committee and the members of the Department of Botany of Columbia Uni-
versity for the excellent facilities provided and for the many courtesies extended
during the meeting.
MISCELLANEOUS BUSINESS
Upon motion the Society voted to appropriate two hundred dollars from any
available funds for use in the support of Phytopathology for 1917.
In response to a request from the Botanical Society of America to nominate a
member of the American Phytopathological Society for the editorial board of the
Botanical Society, Dr. A. G. Johnson was reconmiended by the Council. The
Society approved of the action of the Council.
The Board of Editors made the following recommendations in regard to Phyto-
pathology, and these were approved by the Council :
(1) That no article be accepted which is written in simplified spelling, but that
in the case of words which have two or more forms in good usage strict uniformity
is not required, but the shorter and simpler forms are to be preferred.
(2) That Phytopathology be issued monthly and include five hundred or more
pages during the year, the price of the Journal to members, including dues, to be
four dollars and to subscribers five dollars per year, the increase in price to begin
January 1, 1918.
(3) Space of one-half page or more at the end of long articles is to be used for the
publication of briefer articles or notes in order to avoid wasting space.
(4) It is recommended to members of the Society that they refrain from pub-
lishing original matter in extension publications, weekly news letters, and other
similar publications which are not usually preserved and permanently filed, also
that such references be omitted from the list of literature.
(5) Recommended that authors of phytopathological papers which are pub-
lished in proceedings of academies, horticultural societies, and other publications of
limited distribution, prepare abstracts covering the original matter for publication
in Phytopathology.
Professor J. B. S. Norton presented a plan for ''a standard chart for per cent es-
timates" in regard to injury and conditions of diseased plants. Upon motion this
plan was referred to a Committee, consisting of L. R. Jones, V. B. Stewart, and
H. B. Humphrey, for consideration and report to the Society at its next meeting.
Dr. E. W. Allen, Editor of the Experiment Station Record, in response to the
resolution adopted by the Society at its last meeting requesting that titles of papers
abstracted in the Record be published in full in the original language, stated that
after full consideration of the matter it did not appear practicable to adopt the
proposed change.
Dr. Donald Reddick, Editor of Phytopathology, presented a verbal report
calling attention to some of the matters discussed by the Board as reported above.
U]>on motion the Society directed the Secretary to publish during the year a new
membership list.
The Secretary called attention to the need of prompt notice of change of address
of members, in order to avoid inconvenience and loss of copies of the Journal, and
unnecessary expense in correspondence. C. L. Shear,
Secretary 'Treasurer
REPORT OF MEETING OF THE PACIFIC DIVISION OF TUB
AMERICAN PHYTOPATHOLOGICAL SOCIETY
A meeting of the Pacific Division was held at the University of Caiifomia, Berke-
ley, December 28 and 29, 1916. President J. T. Barrett, of the CitruB EzperimeBi
Station, Riverside, California, presided at the sessions, while Ralph E. Smith, of
Berkeley, California, acted as secretary in the absence of W. T. Home, of Berkeley
who is spending a leave of absence in Cuba. At the business session of the Society
the following officers were elected for the coming year:
President f H. P. Barss, Corvallis, Oregon.
Vice President, James McM urphy, Leland Stanford Junior University, PkJo Alto»
California.
Secretary-Treasurer, W. T. Home, Berkeley, California.
The following papers were presented:
Apple rosette. M. A. Willis
No abstract.
^11 Alternaria blight of tomatoes in California. Bruce Douolas
No alMtract.
Sour rot of lemons. Clayton O. Smith
No alMtract.
Stem-end decay of Valencia oranges in transit. Clatton O. Smith
No abstract.
Some effects of sulphur on soils. H. S. Reed
Sulphur in the form of elemental sulphur, sulphides, or sulphates is widely used
as a fungicid«*. Much of thiH material finds its way eventually into the soil. Itm
eflfert as a soil constituent is thrn*fon* pertinent.
Under anaerobic conditions microorganisms may reduce sulphates to sulphites
or 8uIphi<ieH. l>oth of which are toxic to vegetation. Oxidation processes may cob*
vert sulphides and elemental sulphur to sulphates. The process is largely, if not en*
tirely. due to biological agencies. If the oxidation process is incomplete sulphite*
may \n* formed.
The hannful effects of sulphur are mon> common in soil deficient in organic mat-
ter, or in MoilH having an acid reaction.
Black-heart dinea^e of the apricot. IIelkn Czar.vecki
StudifJi on Monilia. Kditii Philliph
No alwtract.
MinctUnncoiu obnervations. Jamea .NfcMrRPiiT
No nl»Htract.
Thr t j/w rimrntal incfstigation of alleged smelter smoke injury in CalattroM County,
i'tllxittrfiUl. W. W. TUOMAS
N«> al»j»tr:irt.
l*Uth\itc%fnt\* rtlated to Phytophthora. J. T. Bakrett
Kor Mime tiiiif three MtrainH, |M>rlui(M diMtinct species, of a fungus, whoee asexual
stAg«*»« rcTM-iiiblf vi-ry rloHely thow of l^ythiacyntis citrophthora Smith dk Smith, have
bet^n uricifr oliservation. Thest* stmins differ mainly from the latter funfue in that
1917] Berkeley Meeting of Pacific Division 151
they produce in culture oospores while the perfect stage has not 3ret been reported
from any culture of P. citrophthora isolated from any variety or species of Citrus.
Of the three strains mentioned, one was isolated from decaying apples in March
1908 in Illinois; one from bark of a young apricot tree in March 1916 in California
and the third from bark of an avocado tree by H. S. Fawcett in May 1914.
A comparison of the three forms with four species of Ph3rtophthora has revealed
a very close similarity of the oogonia, oospores, and antheridia to those of P. cactorum
(Cohn& Leb.) Schroeter, while their asexual spores, (sporangia and conidia) differ
mainly in the manner of separating from the hyphae.
This marked similarity of their sexual organs to those of P. caciarumf and of
Iheir sporangia to those of Pythiacystia citrophthora would seem to indicate a close
relationship of the two genera Pythiacystis and Phytophthora.
VaricUiona in Colletotrichum glcBoaporioidea. O. F. Burger
Cultures of Colletotrichum glceosporioides were isolated from different Citrus spe-
cies in California. They have been grown on six different media and each strain re-
sponds differently to the media. The size of the spore, depends in part upon the
medium used. A hundred spores of each strain were measured and it was found
that the mean spore-length of most of the strains, when grown on green bean pods,
is 15 microns. Other strains were found, however, which have a mean spore-length
of 12 and 17 microns respectively.
The cultures can be classified according to their mycelial characters, when grown
on artificial media. Class A. Mycelium dark, olive color, giving a fluffy growth with
but scant spore production. Class B. Mycelium dark, appressed but abundant
spore production. Class C. .White mycelium and abundant spore production.
Sexuality in Cunninghamella. O. F. Burger
Pure cultures were made from single spore heads of Cunninghamella hertholletia
and during the entire work no zygospores were formed in the culture tubes. But
when two strains, whose gametes were compatible are contrasted in an agar plate
zygospores are produced at a point where the cultures meet.
A sexual reaction did not occur with Blakeslee's Mucor V, plus and minus, or
his plus and minus strains of Cunninghamella echinulaia. The strains of C. her-
thoUetiae which acted as neutrals with these two fungi formed normal zygospores
when contrasted among themselves.
The peculiarity in their method of conjugation is, that there were marked differ-
ences in their ability to conjugate with certain strains. Strain A will cnjugate
with strains B and C and strains B and C will also conjugate and form normal zy-
gospores. Cunninghamella bertholletias is therefore a pseudo-he terothallic mucor.
Curly (op of the sugar heel. Ralph E. Smith
No abstract.
One session was devoted to a discussion of the so-called non-parasitic or physio-
logical plant diseases, attention being paid to a number of obscure troubles which
are of particular importance in the Far West. A discussion also took place concern-
ing possible means of making the Society more useful and securing a representative
attendance from the different states at its meetings. The acting secretary was in-
structed to take up the latter subject with members in the territory covered by the
Division to see what can be done along this line.
W. T. HORNE,
Secretary 'Treasurer
LITERATURE ON AMERICAN PLANT DISEASES'
Compiled bt Eunice R. Oberlt, Librarian, Bureau op Plant Industrt and
Florence P. Smith, Assistant
December, 1916, to January, 1917
Alaska Afiicultttral Ezperlmant Stations. Plant diseases. Alaska A^r. Ezpt.
StS8. Rpt. 1915: 39-41. 1916.
Allard, Harry Ardell. A specific mosaic disease in Nicotiana viscosum dialiiiei
from the mosaic disease of tobacco. Jour. Afcr. Research 7, no. 11:481-486,
pi. 35. December 11, 1916.
Avema Sacci, Rosarlo. Molostias cryptoKamicas do cafdeiro. Bol. Agr. [Sic
Paulo) 17, no. 10: 790-840, fig. 157-285. Outubro, 1916.
Baker, Carl Fuller. Additional notes on Philippine plant diseases. Phil. Agr.
and Forester 6, no. 3: 73-78. July, 1916.
Bancroft, C. Keith. Report on the South American leaf disease of the Para nil^
ber tree. Jour. Bd. Agr. Brit. Guiana 10, no. 1 : 13-33. October, 1016.
Funicladium macronpcrum; die back.
Boyce, J. S. Pyrnia of Cronartium pyriformc. Phytopathology 6, no. 6: 446-447.
December, 1916.
On PinuM ponder ima.
Brandes, E. W. Report of the plant pathologist. Porto Rico Agr. Expt. Sta.
Rpt. 1916: 34 35. 1916.
Kxperiments in the control of a banana disease; miscellaneous experiments.
Brooks, Charles, and Cooley, Jacquelin Smith. Temperature relations of apple*
rot fungi. Jour. Agr. Research 8, no. 4: 139-164, 25 fig., 3 pi. January 23,
1917.
Ckivers, Arthur Houston. .\n epidemic of nist on mint. Mycologia 9| no. L: 41-
42. Januar>', 1917.
/'iirntnio Menthtr Pcrs.
Cook, MeMlle Thurston, and Wilson, Guy West. The influence of the^tannin
content of the host plant on Endothia parasitica and related species. New
Jomey Agr. Fbcpt. Sta. Bui. 291. 47 p. 1916.
Short<T article with same title published in Bot. Gas., November, 1915.
Not4*d in previous lif«t.
Dodge, Bernard OgiMe, and Adams, James Fowler. Notes relating to the Gymno-
nfMirangi.i on Myrira and (\imptonia. .Mycologia 9, no. 1:23-29, 1 fig., 3 pi.
Jiiiiuarv. 1017.
1 I'hM list aim* to iDclu'l** tb« i»u)ilu-atioo« of North an*! South Amoriea. th« WttI ladia Mnadn. 9m4
laias'ls rootrol»»l b> tho Uait«ii 8t*tM. aa<l mrtielM by Amariraa WTit«n apitoMiag in foralp
All author* art urg*d to rodpcrat* ia makinc iho liat i-onpWta b> Madiat thoir ■tiiaiiitM wad bjr
iBc rorr«> tiooaaal a-l-iitioaa. ao-i ••pocialU h\ ralltai attoatioa to m«ritoriou« artirlct pnhliillil
of r«cul*' lournala ll«i»nBU or corT«aioaii«ar« ahouM t»« a*MrMM«i to Mmm E. R. Ulicrly, UbtSffteB.
HuTMu of Plant In luairy. U. 8. IHpt. Apir., Waahiagtoa. D. C.
1917] Literature on American Plant Diseases 153
Blliott, John A. The sweet potato ''soil rot'' or ''pox'' organism. Science n. s.
U, no. 1142:7(»-710. November 17, 1916.
Cyaiospora batata gen. no v., sp. nov.
Fawcett, George L. A Porto Rican disease of bananas. Porto Hioo Agr. Expt.
Sta. Rpt. 1916: 36^1. 1916.
Closely resembles Panama disease.
Fawcett, Howard S. Citrus scab. Phytopathology 6, no. 6: 442-445. December,
1916.
Discussion of article by Grossenbacher.
A Pythiacystis on avocado trees. Phytopathology 6, no. 6: 433-435. De-
cember, 1916.
Fromme, Fred Denton, and Thomas, H. E. The root-rot disease of the apple in
Virginia. Science n. s. 46, no. 1152: 93. January 26, 1917.
Xylaria sp.
Germano de Souza, Raul. O Tylenchus acuto-acudatus ('i ) nos cafezaes de Indaia-
tuba. Bol. Agr. [Sfto Paulo] 17, no. 9: 726-736, 7 fig. Setembro, 1916.
Gile, Philip Lindsey. Chlorosis *of pineapples induced by manganese and car-
bonate of lime. Science n. s. 44, no. 1146: 855-857. December 15, 1916.
Gfissow, Hans Theodor. Canada's white pine possessions are threatened with ex-
termination. An authoritative discussion of whit« pine blister rust. Canad.
Forestry Jour. 18, no. 1 : 900-906, illus. January, 1917.
Leaf roll in tomatoes? Phytopathology 6, no. 6:447. December, 1916.
Hartley, Carl, and Pierce, Roy Gifford. The control of damping-off of coniferous
seedlings. U. S. Dept. Agr. Bui. 453, 32 p., 1 fig., 2 pi. 1917.
Pythium debaryanum; Fusarium monxliforme; Corticium vagum var, Solani.
Heald, Frederick De Forest. Some new facts concerning wheat smut. Proc.
Washington State Grain Growers, Shippers and Millers Assoc. 10th Ann. Meet-
ing, 1916: 38-45, 2 fig. 1916.
Huard, Victor Am6d6e. Les principales espdces d'insectes nuisibles et de maladies
v^g^tales. [Quebec Dept. Agr.] Bui. 23, 75 p., 78 fig. 1916.
Hubert, Ernest B. Celluloid cylinders for inoculation chambers. Phytopathology
6, no. 6: 447-450, 1 fig. December, 1916.
Jehle, Robert Andrew. Means of identifying citrus canker. Quart. Bui. State
Plant Bd. Florida 1, no. 1: 2-10, 12 pi. (partly col.). October, 1916.
Johnson, James. Enfermedades del tabaco y manera de combatirlas. Parte V.
Hacienda 12, no. 4: 124-126, illus. Enero, 1917.
Johnston, John Robert. Enfermedad de la raiz de la cafia de azticar. Hacienda
12, no. 4: 117-118, iUus. Enero, 1917.
To be continued.
Marasmius on sugar cane. Mycologia 8, no. 2: 115. March, 1916.
Comparison of M. stenophyllus and M. 8<icchari.
Lewis, A. C. Cotton wilt in Georgia. Georgia State Bd. Ent. Bui. 40, 18 p., 8
pi. 1915.
Varieties of wilt resistant cotton, p. 8 ff.
McClintock, James A. Is cucumber mosaic carried by seed'i Science n. s. 44, no.
1144:786-787. December 1, 1916.
Peanut mosaic. Science n. s. 46, no. 1150: 47-48. January 12, 1917.
Sclerotinia libertiana on snap beans. Phytopathology 6, no. 6: 436-441,
2 fig. December, 1916.
McCobbin, W. A. The white pine blister rust: does the fungus winter on the cur-
rant? Science n. s. 46, no. 1152: 87. January 26, 1917.
154 Phytopatholoot [Voi- 7
Martin, WillUm H. Influence of Bordeaux mixture on the rates of trampirmtion
from abeciflcd leaves and from potted plants. Jour. Agr. Research 7, no 12:
52^-548. December 18, 1916.
Literature cited, p. 547-548.
Maatey, L. M. The hard rot disease of gladiolus. New York Cornell Agr. Expi,
Sta. Bui. 380: 149-181, flg. 38-14, pi. 15^16. 1916.
Bibliography, p. 180-181.
StpUnia Gladioli,
Mandlola, Nemeslo, and Bspino, Rafael B. Some phycomycetous diaeaaea of cul-
tivated plants in the Philippines. Phil. Agr. and Forester. I, no. 3: 65-71,
illus. July. 1916.
Bibliography, p. 71.
Phytophthora diseases; Pythium Ekbaryanum.
Mix, A. J. Cork, drouth spot and related diseases of the apple. New York State
Agr. Expt. Sta. Bui. 426: 473-522, 12 pi. 1916.
Nonparasitic.
' Sun-scald of fruit trees, a type of winter injury. New York Cornell A^.
Expt. Sta. Bui. 382: 233-284, fig. 60^1, pi. 18-19. 1916.
Literature cited, p. 283-284.
Probably a winter injury caused by direct freesing to death of the tiaaue.
Morse, Warner Jackson. Studies upon the blackleg disease of the potato, with
special reference to the relationship of the causal organisms. Jour. Agr. Re-
search 8, no. 3: 79-126. January 15, 1917.
Literature cited, p. 124-126.
Hanllu* atronepticun Van Hall.
Nowell, William. Fungoid and bacterial diseases. West Indian Bui. If, no. 2:
133-143, 146 147. 1915.
Listed under host and locality.
See also p. 121-122.
Fungous and bacterial diMoases. West Indian Bui. 16, no. 1: 17-25,
1916.
Listed under host and locality.
See also p. 2.
Hosellinia root diMMifles in the I^esser .Antilles. West Indian Bui. li, no.
1:31 71. 12 fig. on 4 pi. 1916.
Keferenc«»H, p. (H) 71.
Orton, Clayton Roberts. Phytophthora infestans on tomatoes in .\ustralia. Phy-
topathology 6, no. 6: 447. Drci'mln^r, 1916.
Osner, George A. I^>af smut of timothy. New York Cornell Agr. Kxpt. Sta. BuL
,3S!: is:> 230, fig. 4:> .W, pi. 17. 1916.
Bihli«»graphy, p. 22«'> 2.10.
f ' •» / 1 higii ntritrform if .
Paul, B. H. Th«* pine blinUT niMt. (\»nwrv. Com. State New York BuL 15, 18 p.,
illiiM.. 1 foM map. [1917.1
Seaver, Fred Jay. Sonu* p:i|N>rH pn>(M*iit4;d during convocation week. Mycolofpa
9, no, 1:42 -U*. Jaiiuiiry. 1917.
Joint fW'Minii of Hot. Sor. an<l Phytopath. Soc, P>iday, I>ecember 28, 1916,
in connection with .\nirr. .\.Hf«i)c. .\dv. S4m.
Smith, Erwln Frink. .Mrrhani^ni of tumor growth in crowngall. Jour. Agr. He-
m-arch 8, no. r»: 1(W> 1S(>. pi. 4 <m. January 29, 1917.
Literature cited, p. is^lW.
|PnYTOPATHoi/)<iV, foF Fehruafv, 1917 (7: \~M, PI. I) wan iiisued Fob-
nmr\' 7, HUT. I
PHYTOPATHOLOGY
VOLUME VII NUMBER 3
JUNE, 1917
STUDIES ON BACTERIUM SOLANACEARUM'
E. E. Stanford and F. A. Wolf
With One Figure in the Text
Studies on the wilt diseases caused by Bad, solanacearum have been
in progress at the North Carolina Agricultural Experiment Station since
1903. A recent bulletin (4) dealing primarily with remedial and pallia-
tive measures for tobacco wilt contains the results of certain of these in-
vestigations. In the present paper are presented data bearing (1) on
the distribution withiil North Carolina of the disease on tobacco {Nicotiana
iabacum) and tomato (Lycopersicon escuUntum), (2) on cultural studies
to determine the identity of the strains from the several hosts and (3)
on the results of cross inoculations.
distribution of BACT. solanacearum within north CAROLINA
Tobacco wilt has probably been present within the state for at east
twenty-five years, but occurs in only a small proportion of the lands de-
voted to tobacco culture. The disease was first definitely recognized
in the southern portion of Granville county in 1902. Records indicate
that the disease has yearly continued to spread so that it now occurs in
eleven counties, namely: Granville, Durham, Vance, Wake, Franklin,
Ashe, Guilford, Greene, Davidson, Chatham and Yadkin.
The losses in the first four of these counties are confined to the southern
portions of Granville and Vance and the northern portions of the adjacent
counties of Durham and Wake. In this area the disease is so generally
destructive that the growing of tobacco has been abandoned on many
farms. The disease is confined, however, to relatively small areas in
the seven remaining counties.
* Acknowledgment is hereby made to the several members of the Department
of Botany and Plant Pathology of North Carolina Agricultural College and Experi-
ment Station, who, since the inception of these investigations, have aided in the
work.
1S6 Phytopatholooy [Vou 7
It is tt striking fart that collections of Imcterial wilt of tomato which
have been vcr>' Rcncrally made throughout the State show that the diaeaap
on tomatoes orcura in thirty-nine counties of the state. It is realised,
of course, that the disease on Itoth tobacco and tomat^x^ may not have
been reported from all localities in which it may occur, yet the fact remsiiu
that the disease on the two host« is not coextensive. No a<lequatc exfdana-
Piii. 1. Map op North Caholika Shiiwino the Known Dihtubltion or
BArTERIVM HOLASArEAKl'H ON" ToSACCO ASO-OS ToMATO '
tion is ut hnnd (o iiccount for this, in xiew of the fact that reriproeal inoni-
latioat nre ho caHJiy i-fTocted and that the strains of Bad. iioiaTiactarum
from IiiIhicco and toniattx-s arv. identical as shown by the cultural atudim
of scvend investifiiitorx and ronfimwd by studies to l»e presented later
in this [uipiT.
It nuKht Ik> ad<lc<l, that no <luta have accumulat^l relative to the di»-
tribulitm of tliis purHHite on otht-r solanat-eous hotts and that the tliseaa»
on peanut (ATtickin hypog(ra) has Ixfn ol>serve<l in one locality only.
nXTIHAL HTrniES
In view of the fact that in wrtain sections. Bad. Kilanacearum ts not
present on ull of the hosln which un- known to l>e subject to attack, a aludy
by means of artiliml cultun-s w:is nmde to iletermine the identity of
the stniinK from the more comnKin hosts. Several investigatots, anioiiK
wbnni iii:iy Is- nientionetl Smith ( '> an<l 0) and Honing (6)' have ptvnouiJy
r(']M>rlvd cxU'iiMve .stu<li<-s on Hact. solaniteearum in artificial culture and
n-giinl t) rgHniMii from (be s«'viTid hosts as identical. Nevcrthplesn.
t Hi-ri-n-nri-H III rill- imMii'iiliiniH of lloninii. HunK<-r. und I'yrdk mi* given in Vol*
uiiK- :l 'if .><iiiil>i'i' tlrK'iirin in Hi-liilion to I'tiinl Diwhwh. Tho tulhora haw not
■rfii l)ir>>^ i>iililii'Bti»iii>. I>iil only ilii' alMtnirtH lavrti \>y I>r. Smith. BiMiocr«phitw
of Hart. ...^l.,.l,■■.,r..n. on p. -.'Is iWi iind 1*70-271.
1917] Stanford and Wolf: Bacterium solanacearum 167
parallel cultures of the organism isolated from tobacco, potato, (Solanum
tuberosum) tomato, eggplant (Solanum mslongena), peanut, ragweed
{Ambrosia artemisiifolia) and Edipta alba have been made. These isola-
tions were made from various parts of the hosts and from plants from several
localities within the state. In general, the cultural variations which
appeared between the strains from the different hosts were no greater
than those from strains all of which came from the same host. Since,
however, certain additional facts are brought to light, a brief account
of the growth in culture is pertinent and is herewith given.
Colonies on agar plates usually become visible within thirty-six to forty-
eight hours at 28°C. At a temperature 4 to 5° lower, they may be scarcely
noticeable at the end of seventy-two hours. They are at first white,
wet-shining and opalescent, circular in outline, slightly raised and with
smooth margin. Colonies become 3 to 5 mm. in diameter in five to six
days. They soon become distinctly opalescent-blue by transmitted light,
when the plates are held some distance from a poorly Ughted reflecting
surface. When the colonies are viewed with the blue sky for a back-
ground, they are distinctly brown with faint concentric rings. Later,
they become very markedly brown by reflected light. The pigmentation
originates at the center of the colony, spreads toward the margin and is
diffused somewhat into the agar. The color is more intense at the cen-
ter of the colony. Subsurface colonies are globose or lenticular, much
smaller than the surface colonies and become brown much sooner than
do the surface colonies.
The growth on agar slants develops slowly, is filiform at first and usually
spreading at the bottom of the slant. It is shghtly elevated and the mar-
IB^ is entire or wavy. Colors and pigmentation develop similar to those
in plate colonies with considerable variation in time of appearance and
intensity of pigmentation which may vary from scarcely perceptible to
brownish black. When Witte's peptone is used, a deeper pigmentation
develops than when Difco peptone is employed. When the colonies
become blackened, the organism is no longer viable. It appears to re- |
tain its vitaUty for about four and one-half months on agar but rapidly '
loses its virulence on this medium.
The appearance of colonies on Utmus lactose agar slants is similar to
those on nutrient agar. The Utmus is at length sUghtly reduced. Pig-
mentation is first noticeable in three to four weeks and a brown stain soon
becomes diffused throughout the agar and masks its color. Portions of
the substratiun which are not brown become sky-blue by diffuse Ught
and reddish plum-colored by transmitted Ught.
The appearance of this organism in bouillon cultures is somewhat
variable. A rather uniform clouding develops within twenty-four hours
158 PHYTOPATHOLOCiY (VoL. 7
which l)erome8 so intenflely opaque within a week as to make it impo»-
nihle to see through a l)Ouillon tulx; when the tul)e is placed immeclistely
in front of an object. Numerous pseudogloeae may appear forming
flocculent particles on the surface. No pellicle nor ring is formed, but
a thin, opalescent scum appears on the surface. A dirty white, viMom
precipitate develops within a week or two. Within four to six weeks,
this precipitate will have l)ecome dense and the supernatant l)ouiIlon
will have liecome cl(»ar at which time the organism Ls no longer \'iable.
Various degrees of pigmentation may occur, l>eing more intense with
Witte's than with Difco peptone. Tul)es of the l>ouillon made with the
fonner l>ecome hn)wnish black on long standing.
(>n potato plugs, the growth is spreading, thin or slightly raised, white
or flc^sh-i'olorefl at first l)Ut rapidly In^comes bn)wn, often pitch-black.
The surrounding licjuid U'coines clouded and bn)wn. The organism is
short -live<l on this n)edium and conuiionly loses its vitality within a
wec»k. The vinil(»n<'e of linct. siflanaccfirum on artificial me<lia is liest
retained on |X)tat() phigs. but transfers must Ik* made at inter\'als of aliout
two davs.
When gn)wn on milk, there is no peptonization nor precipitation of
ca.«ein. A slight visci<lity and the odor of putn»faction are developed
and the medium at length In^comes bn>wn and alkaline.
The organism (*auses n partial clarification of litmus milk with a ck^p-
ening of th<» blur color which apfx^ars n*ddLsh by transmitted light. A
slight dirty white i)n»cipitate is fonned, which lK»(*i>mes brown on loiiK
standing. The ba<*teria may n»main viable* for five months on this me<lium.
The surface coloni<»s on gelatin are small, circular, white and wet-shin-
ing. Submerge<i colonies an* glolMist* and yellowi.sh to brownish, (irowth
aU»ng the line of the stab on gelatin is white, later l)ecoming bn)wn, fila-
mentous and lH*st at the surfacr of the medium. No liquefaction cMM'urK.
TIm'H* is no evident gn)wth on Crohn's .solution in four weeks. C>n
I'schinsky's soluti<»n, growth ranges fnun none to feeble with slight
clouding.
Ill (lrxtn>s«» bn»th. a copious gn»wth develops in the opi»n arm, extc^nd-
ing (»iily to thr bas4' of the* <*Ios4mI ann. .\n abundant, rather viseoiis
|)rf('if)itatr ap|M>ars. Th<> nHMlium at Irngtii lK*comc*s bn)wn in the o\H*n
ann and i^ strongly alkalinr with no evolution of gits.
< ultural <*harartrrs on s;i<*charos<* bn>th an* similar to those* on tlextrnwe
with a l«»s> marked tend«*nrv to the development of a bmwn color.
(Irn\%th i*« fe<*ble on la<ioM* brcith with little .s<m I i mentation and little
or ni» bniwii eolur i?^ devel<»|MMl, even after s<»ven w<»<»ks. The acidity of
aril! broth is dimiiiishe<{ l>ut neutral broths an* not rendered alkalin.
<>n niannit. the growth eharaeters an* similar to thos4» on lactos<\ .\ ver>*
1917] Stanford and Wolf: Bacterium solanacearum 159
copious growth ensues in glycerin broth with a marked development of
brown color. Growth on maltose is similar to that on dextrose. Nitrates
and ammonia are formed in moderate amount in nitrate solutions.
Growth is much delayed and diminished in hydrochloric acid +25
Fuller's scale, and is entirely inhibited at +30. No growth occurs in
double strength bouillon rendered +33 acid by the addition of expressed
tomato fruit juice. Smith (6) reports growth in +33 acid of beef juice.
The optimiun reaction lies between +10 and +15. ^
Growth is slight or none in bouillon --5 with sodium hydroxid. No
growth occurred in —10 sodium hydroxid. The organism is little re-
tentive of vitality on culture media. Milk appears to be the best medium
for long continued growth on Bad, solanacearum. The organism may
remain viable for two months in sterilized distilled water. No evidence
of diastatic activity was found when the organism was grown on potato
plugs.
A considerable number of special media have been prepared, among
which are soil extract, casein agar, Hey den's Nahrstoff agar, potato agar,
potato leaf agar, and tomato leaf agar. No growth of diagnostic signifi-
cance developed on any of these media.
Bacterium solanacearum is very short-lived in mixed cultures. Honing
(6) noted a marked antibiosis between the wilt organism and B. mesen-
tericus as well as other species plated from wilted tobacco. In our studies
also, various bacteria have been found to replace Bad, solanacearum in
decaying, wilted plants. Five strains of yellow chromogens isolated
from diseased tomatoes, tobacco and peanuts were found in the fall of
1915 to exhibit marked antagonism to the wilt organism. In inter-
secting streaks on agar plates, the chromogens tended to crowd out the
parasite. The presence of B, mesentericus and other soil inhabitants
appear never to be so antagonistic, however, as to eliminate Bad, solanace-
arum from infested soils.
Thus far attempts to isolate the parasite directly from infested soils
have been unsuccessful. This is due in part at least to the fact that
Bad, solanacearum is inhibited by other soil inhabitants which develop
on the plates. Honing (6) however, succeeded in isolating it on plates
from dilution cultures of well water.
cross-inoculation experiments
Bacterium solanacearum has previously been shown to attack mem-
bers of eight widely separated families, Urticacese, Leguminosae, Tropaeo-
lacese, Euphorbiaeeae, Verbenaceae, Solanaceae, PedeUaceae and Compositse.
The organism was first described by Erwin F. Smith (5) in 1896 as
160 Phytopathology [Vol, 7
the cauae of a wilt disease of tomato, eggplant and potato and he succesa-
fully inoculated Solanum nigrum, Datura stramonium, D. mdelUndeM,
D. foMuosa, D, cornucopia^ Physalis crasnfolia, P. philaddphiea and
Petunia (hybiid).
Several investigators, among whom are Hunger (6), Stevens and Sackett
(7) and Uyeda (6) have reported a wilt disease of tobacco. Honing (6)
in 1910 first reported this organism as the cause of disease in plants out-
side of the fk)lanacea*. He found it in Pouzolzia sp., Phyaalia angulata^
Indigofera arreeta, Arachis hypogceOy Mucuna sp., Acalypha boehmeroideM^
Ageratum conywides, SpUanthes acmella, Pluchea indica, Blumea baUami'
fera, Synedrella nodiflora and Tectona grandia. He also successfully inocu-
lated several ornamental varieties of Nicotiana, Capsicum annuum and
Sesamum orientate,
A wilt disease upon peanut was sul)sequently reported from North
Carolina (2) and later studies (3) in this state added two composites.
Ambrosia artemisiifolia and Eclipta altni, to the list of naturally infected
hosts.
A wilt of nasturtium (Tropetolum maju^) caused by Bad. sotanacearum
was reported from Maryland by Br>'an (1). She succeeded in inoculating
also the common cultivated Agc^ratum and Verbena.
The artificial inoculation ex|x^riments conducted at the North Carolina
Kxperinu*nt Station prior to 1913 were confined primarily to solanaceous
plants. When in the summer of 1912, it was found that peanuts are
subject to attack by Bact, solanacearum, this host was successfully inocu-
lated with strains fn>m tobacco, peppers and peanuts. The strains from
peanuts wen* also found to l)e pathogenic to tobacco.
During the season of 1914, Bact. solanacearum was isolated from di»>
ease<l ragi)^'ee<ls (Ambrosia artemisiifolia) and subsequently found to be
prcKluctive of wilt on tobacco, tomato, potato, Eclipta alba and ganlen
nasturtium (TropuK^lum). The recipr(H*aI inoculations upon ragweed
with strains fn>m tobacco, tomato, potato and Eclipta aU>a were rather
unsucc(*ssful. No systemic invasion n^sulting in death, but merely a
local o<*clusion and blackening of xyl<»m elements occurred in inoculated
plants.
In tin* fall of 1915, strains isolatiNi from wilted Eclipta atba were auc-
c(*Ksfully inocuhit<'<l into tomatoes, potatoes, tobacco, garden nasturtiunui
and Eclipta allnt, A nit her more comprehensive series of inoculations
on rultivatcd an<i wild s|)4*cies was in.stitut4*d in 191G, the results of which
an» hcn»in brit^Hv sum!nanze<l.
Mrthtti of vHticultittifU. The strain of Bact. yolatmcearum emplo^Td in
niakiiig the initial inixulations wa.s (»btaincd by the pounxl plate method,
fnim wilted t4>barro plants from Cn'^Mlmoor. North Carolina. As sooo
1917] Stanford and Wolf: Bacterium solanacearum 161
as the organism had developed on these poured plates, transfers were
made to potato plugs. Rep)eated transfers at intervals of one to three
days were made on this medium. The pigmentation on old cultures on
agar and on potato plugs was regarded as sufficiently characteristic to
estabhsh the identity of the wilt organism. Inoculum from one- to three-
days-old cultures on potato plugs was used in all of the inoculations.
Inoculations were made by pricking the plants near the tips of the branches
and inserting the inoculum. A niunber of check plants, either iminjured
or pricked with a sterilized needle were used in the case of each species
tested. Since it was known that Bad. solancLcearum loses its virulence
even though repeated transfers are made, no attempt was made to use
the original strain from tobacco in all of the inoculations. Instead, isola-
tions from certain of the inoculated species were used in continuing the
series of inoculations. Some differences in virulence appeared in strains
which had passed through different hosts but no such marked decrease
occurred as when the organism is repeatedly transferred on culture media.
In general, young, vigorously growing plants were used in these tests,
although in some cases, rather matiu*e plants were employed. The cul-
tivated species were grown either in the greenhouse or in small experi-
mental plats at West Raleigh, North Carolina and the weeds grew in
waste places where they could be kept under observation for the neces-
sary length of time. In general, as soon as inoculated plants showed
s3anptoms of disease, they were examined microscopicaUy to determine
the presence of bacteria within the tissues at points remote from the point
of inoculation. The organism was then reisolated by the poured plate
method, and its identity established by the characteristic growth on agar
and potato plugs. As supplementary evidence, the reisolated organism-
was inoculated into tomatoes or tobacco.
The accompanying diagram of the plan of these cross-inoculation experi-
ments has been so arranged as to show at once the source of the inoculum,
the result of the inoculation and the number of plants inoculated.
ResuUs. When comparison is made with the host species previously
enumerated, it will be seen from this tabulation of the results of cross
inoculations that the following plants 'have heretofore been unreported
as subject to attack by Bad, solanacearum: Stizolobium niveum, Tropceo-
lum lobbianumy T, peregrinum, Croton glandulosus var. s&ptentrionalia,
Impatiens batsamina, Verbena erinoides, Lycopersicon cerasiforme, L.
pyriformey Browallia demissa, Physalis alkekengij Schizanthus pinnatus,
Salpiglossis sinuaia and Martynia proboscidea. Twelve of these species
belong to families representatives of which had hitherto been known to
be subject to attack and one species, Impaiiens balsaminay belongs to
an additional family.
162 PinTOPATHOLOOY [VoL. 7
I "-si*
"■ B ■• •
? 3 S J
i 7
--5 5
, Jil:
3 « t : p I i
2 S^ 5 • k ?
i J «c - fc " 1
P ^
!.mj
1917] Stanford and Wolf: Bacterium solanacearum 163
It may also be noted from the tabulation that no infection resulted in
Stizolobium niveum and Physalis alkekengi when inoculated with the or-
ganism isolated from wilted Impatiens balsamina. Further, no demon-
strable infection resulted in the case of Petunia (hybrid), Datura cornu-
copia, D, fastv4)8a, and Physalis alkekengi when the isolations were made
from wilted Browallia demissa. Smith (6) had previously shown the
first three of these forms to be subject to attack. Since Stizolobium niveum
wilted when Datura tatula was the source of the inoculum and Physalis
alkekengi, when wilted Verbena erinoides was employed, it is indicated
that virulence is influenced by the host plant.
In the following species, Uttle or no external injury resulted from inocu-
lation, but the vascular tissues were found to be invaded: Euphorbia
nutans, Solanum carolinense, Physalis angulata, Impatiens sultani, Bid^is
bipinnata and Erigeron canadensis.
Inoculated plants of Ambrosia artemisiifolia and Eclipta alba wilted
thus confirming previous studies (3).
In general, it can be said that the external symptoms and pathologi-
cal histology of the plants which were artificially inoculated in these studies
differed in no essential particular from those of other species which have
previously been reported as hosts for Bact. solanacearum. The 8p)ecies
of Tropseolum, Lycopersicon, BrowalUa and Eclipta tested are to be
regarded as very susceptible, whereas, Stizolobium niveum and Physalis
alkekengi appear to be very resistant. That Stizolobium niveum is highly
resistant is shown by the fact that in a field test at Creedmoor, North
Carolina, no demonstrable infection develop)ed in any of the plants grown
in wilt-infested soil. It is interesting to note that when young ragweed
plants grown in the greenhouse were inoculated, they quickly succumbed
to wilt, while numerous individuals grown out of doors when inoculated
with the same strain showed no external symptoms of disease. Little
external evidence of disease developed in rather mature plants of Croton
but young plants were easily wilted. In the case of Impatiens balsamina,
the foliage became slightly wilted, some distortion of the stems occurred
and adventitious roots were formed. The discoloration of the vasculaV
bundles of the stems showed through the cortical tissues as brown streaks.
This species was found to wilt slowly when inoculation was effected by
potting plants in infested soil. The discoloration of the vascular system
is externally visible in wilted stems of Eclipta alba and the leaves become
characteristically crisp and blackened.
The economic bearing of these additional weed and cultivated host
plants for Bact. solanacearum upon the problem of wilt control is at once
apparent when it is indicated that certain of these forms, namely; Erigeron
canadensis, Ambrosia artemisiifolia. Euphorbia nutans, Croton glandulosus.
164 Phytopathology [Vol. 7
and Solanum carolinense are widespread in cultivated fields in the State.
Edipta alba is often found in ill-drained lands. Datura tatuia and Bidenti
bipinnala are not uncommon weeds about farm buildings and lots. The
Lycopersicons and Martynia probascidea are locally rather common in
gardens. The Tropsolmns, Verbena, Impatiens, Browallia, Schixanthus,
Salpiglossis and Physali^ alkekengi are more or less commonly grown as
ornamental plants. The results with velvet beans (Stizolobium niveum)
which is related to Mucuna mentioned by Honing (6) are significant
since this crop is becoming of considerable importance in the South. At
least, it cannot l)o recommended that velvet l)ean8 Ix" grown in a rotation
s\'8tem in soils infested with Bad. aolanacearum.
SUMMARY
1. A wilt of tobacco caused bv Bad, Holanacearum has been observed
in North Carolina in eleven counties and a tomato wilt caused bv the
same organism has U^^^n noted in thirty-nine counties.
2. Pn»viou8 cultural studies on the idcntitv of Bad. solanacearum from
various hosts are (Confirmed since the variations which appeare<l in the
strains from tobacco, (Nitato, tomato, eggplant, fx^anut, ragweed and
Ediftta alba were no gn»atcr than in strains all of which came from the
same host.
3. A new family of phanerogams, Balsaminac(*ip, has been addcnl tii
the nuinlxT previoiu<ly reportcil to contain host six'cic'S of Bad. solana-
cearum. MemlM*rs of nine families arc now known to Ih' subject t4) attack
by this organism. Thirt(M*n additiomil si>ecies of plants, classifieil as
follows showe<l wcll-<h'fincd wilting or serious injur>':
Ii4*ginHinosa* Sticolobium nm^um
Tn)j)aH)lacea» Trofxrolum lobbumum, T. lyeregrinum
Kuphorbiaceu* Croion glandulosus var. neptenirionali^
Balsa in ina(*ca> ImfHitietM balsamina
VcrlH*nu<M*a» \'t'rhi'nn vrifundeH
Soluna(M»a» Lycofpertfmm crrasiformr, L. pyriformr, Brovnil*
I in (i4'mis8a, Physalis alkekengi, SdiisanlhuJt
pinnntuii, Salpiglossis siniiata
VriM'iiivviv .Martynia prttl}Osci(U'a.
No outward si^n** of disease (|rvcl(»|M <1 in the vnsv of six other s|M»cie3*
in whirh the org:uii*<tii nuiltiplied rapidly within the vascular |)ortioas.
Fivf 4»f these sfNMMes an* previously unre|K)rt«'d, mmicly:
Kuphorbiacea* Euphorbia nutans
Solaiuieea' Stdanum carolimnse
Hals^iiiiinaeea' Im/nitiens sulidini
( *on 1 1 ¥ »-« i t a' B idens bipin nata , Erigeron ca tuidensiM.
1917] Stanford and Wolf: Bacterium solanacbarum 165
literature cited
1. Bryan, Mary K. A nasturtium wilt caused by Bacterium solanacearum.
Jour. Agr. Research 6: 451-157, pis. 63-64, figs. 3. 1915.
2. Fulton, H. R., and Winston, J. R. A disease of the peanut caused by Bac-
terium solanacearum. Nofth Carolina Agr. Exp. Sta. Ann. Rept. 86 and
87: 4a-47, figs. 4. 1913-14.
3. Fulton, H. R. and Stanford, E. E. Two wilt hosts of Bacterium solanacearum,
Phytopath. 6: 108. 1916.
4. Garner, W. W., Wolf, F. A., and Moss, E. G. Control of tobacco wilt in the
flue-cured district. United States Bur. Plant Ind. Bui. (In press.)
5. Smith, Erwin F. A bacterial disease of the tomato, eggplant, and Irish potato,
(Bacillus solanacearum n. sp.) U. S. Dept. Agr., Div. Veg. Phys. and
Path. Bui. 12: 1-28, ph. 2. 1896.
6. Smith, Erwin F. Bacteria in relation to plant diseases. 8: p. 309, pU. 47, figs.
155. 1914. Carnegie Institute of Washington, D. C. Publ. 27.
7. Stevens, F. L. and Sackett, W. G. The Granville tobacco wilt. North Caro-
lina Agr. Exp. Sta. Bui. 188: 81-96, figs. 16. 1903.
SPARASSIS KADICATA, AX rXDKSCKIBKD FUN(;US ON THK
HOOTS OF CONIFERS
j a m k h u. \v k i r
With Fivk Fic;!kks in tiik Tkxt
In August. 1912, the writer collected s(»vt»nil siMTiinens of a sp<M'i<'> <>f
Sjwirjissis growing on th(» nnits of various (•onif(»rs in the Priest River
Valley, Maho. Lloyd, to whom sjMM'iniens wen» sent, pronounced il an
inuh»scrilKMl sjx'cies. Cotton of the Pathological Laboratory* at Kew wlio
was a<lvis4M| hy Lloyd of th<» writ<*r's six'ciniens stated that the plant
w:ts unknown to him. Since collecting the first s|H»cimens. the writer
luis studied the |)lant in several regions of the Northwest and finds tlutt
of the many jx^culiarities of the s|KM'ies the most surprising diM*ovt»r>"
is its evi<lent parasitism on the nM)ts of conif<Ts. Although this fact was*
noted in MM 2. it was not until the plant was carefully studied in ix> n-b-
tion to its S4»V(Tal ho>ts that this ph:ise in its life historv eouhl Ih* snti*-
factorilv determined.
nKS<'HIPTIO\ OF THK KrNCiT.S
Since the fungus d<M»s not agn»<» with any known inemlnT of the genu^.
it is described as new.
Sparassis radicata n. sp.
Fruiting stnictun* larg(^ 12 to 22 cm. ))road. 10 to 1(> cm. high. dilat«-<i
aUive. compact. Heshy. tough, whitish, cn^amy yeUow with age. I>ninclif«|;
hninches numerous, hori/^ontal or vertical, anastomosing, sometimes fonn*
ing lahyrinth-like cavities. nion» often comiKictly arnmgiMl. ver>* thin, fan-
shafMMl with wavy, sometimes din-ply IoImmI margins, ocnt.'iioniilly striat«»<i.
aiiiphigcnou*^ or unilateral, de|M*nding on the fMKsition of the l>nin<*h.
Malk. H'lrrotioid. tuU'rculate, firm, solid, sometimes hnincheil, 21) ^{l> mi.
long. .'» s rm. hroail: s|)on*«i, ..">()< nmgc 2.S 4.0 X 2.S-r>..') ^, standard ^ir.9*
'.iV .' '».l ^. oviiid. hyaline.
7*v/*» UH-nUtij, Pri<»st River. Idaho
Ihilntni. Living kmiI-^ in c'oiiiferous tn'4'^'.
litiHtj* . nn*Kon. Idaho. \Va.*«hington. Montana, and British (\»luml*ia.
T\tlH- tniit*rt'il. In the nffin* of Inv("«tigatio!w in Forest PatholiH^y,
liun-riu fif Plant Industry. Mis.s^nila, Mont.
1917] Weir: Sparassis radicata 167
general morphology and taxonomy of the genus sparassis
The genus Sparassis was established by Fries* and placed in the Cla-
variaceae because of its frondose habit, fleshy consistency, and the be-
lief that the spores were produced on all surfaces of the sporophores. It
has recently been shown by Cotton^ that the hymenium of Sparassis is
not amphigenous but that the flattened branches with the exception of
those in the center of the sporophore, are unilateral. On the basis of the
flattened sporophore and the inferior hymenium Cotton suggests that
Sparassis should be removed from the Clavariaceae and placed in the
Thelephoracea*. He points out that in the Merisma section of the genus
Thelephora are species with upright, partly unilateral sporophores either
terrestrial or growing on wood which in many respects have the charac-
ters of Sparassis. In points of smoothness of the hymenium he further
suggests that Sparassis is alHed to Stereum but since the relationship
to Stereum is not very close sees no reason why the genus Sparassis should
not be transferred to the Thelephoracese without reference to any par-
ticular genus. Sparassis would then be distinguished as a genus of the
Thelephoracese having fleshy, flattened, horizontal or vertical anastomos-
ing branches with unilateral structures. The same view is entertained
by Lloyd' who thinks the definition as laid down by Fries *' fertile on
both sides ^* should be corrected. Whether or not this view should be
adopted in view of the fact that there is considerable irregularity in the
formation of a unilateral sporophore is doubtful. In young sporophores
of Sparassis crispa (Wulf.) Fr. examined by the writer, also of Sparassis
radicata^ the hymenium is by no means confined to the lowermost portion
of the flattened branches but is found more or less uniformly over all free
surfaces. This is particularly true, as Cotton points out, for those
branches in the center of the sporophore but with a more pronounced
unilateral structure toward the periphery. The hymenium of Sparassis
radicata is formed very rapidly on the reverse side of the peripheral lobes
when changed from their origninal position. A few specimens with un-
usually vertical lobes showed an amphigenous hymenium throughout
making it seem probable that the lobes only become unilateral when they
develop in a position allowing the influence of gravity to be more active
on one side than another. There are, however, very few unilateral fungi,
if any, that, under proper conditions of growth, will not when reversed
develop the hymenium on the upper side.
' Fries. Systema mycol., I, p. 462.
• Cotton, A. D. On the structure and systematic position of Sparassis. Trans.
British Myc. Soc. 1911: 336-339.
» Lloyd, C. G. Letter No. 61, note 400.
168 Phytopatholoot [Vol
Until QiH'Ict* nn<l PatouilInnl* iiotni the affiiiiticx of Sparassis with ci
tiiiii ifroiips in l)ii' Thc-lt-pliomrcii' Imt iippamilly without ilcfinilr knot
i-djEi' of ilic liyiiirnial ilfvclopiiiciit in llic (jt'iiiis. Somi-wliat later Mai
M-iuinilcit Spitrassis fniiii tin- < 'lavariiH-i'u' making it the ty|>p «f a «pw
family, tlic SiMirassiili>ii>. AllhoUKli Muirc's (-lassifii-alion wiis udoptini
l^itsy,' rritioal work on thi* licvi-loiinicnl of the hyincniiim. ixTmanpn
"Iftlk :iltKrlin) Ui rhr root*
nf ih.. .iMihl.-ral -ini.-l.in- <,f ih.- l,raiu-li<.. niul.-r vari.ni^ fa.-l..r^ ..f rh.w
aitil -.. lortli i- wry i.iu.l. ii.-.-l.-.i Ui,,r.- tli- .■ImiiKi- .-li.aiM U' a.-r.-pf
l.l..y.l La- mail., ll..- xiKU.-rnm t,. tli.. wnt.., lli:,t rli.. u.-iiii- Sp^.ra-
f:tll- naliirally iiili> Iw.. M-.-ljnii-.: tii-l >', r<,v,,„, ii|„.|| „ „i,.ir ».f ;, n.-l
' l^i.-[.i Kl..r:. [i.vr..|<.Ki<|ii'- 'i>- l;i KriM.'.. I'urj*. Ivvs.
' I': ill^r.l \j~ llviix'i yW'l.'. .i'Kiir..]- I'urU IS.S7.
■ \I:,.r. ItirK-r. Iir- .-> l..l..Bi.|ii. - .r l,.\..Mm,ii.|i.,> -ur 1.-. H«-i.l..rnv.-f.tf ,. A
iu-\.- :,<t Hull S.M' iinr, tr:iii.r. 18: -, I'.Nrj.
' I...t-> V..rir.p' iil-r l-.tuiii«li.- Si:,i ,. (pmliLilnr. t. Jrna. lUttV
1917] Weir: Sparassis radicata 169
nature especially near the base of the branches and is very closely re-
lated to Clavaria, and a second section consisting of species with thin
lobes such as S. spathulata in the United States, S. laminosa of Europe,
and S. radicata which has thinner lobes than either of the former. This
seems to be a very logical arrangement. He further suggests^ that Spa-
rassis laminosd and S. spathulata are probably indentical. Their sporo-
phores are certainly very similar.
THE ROOT STALK OF SPARASSIS RADICATA
Sparassis radicata (fig. 1) differs chiefly from Sparassis crispa(WuU.) Fr.
{S. ramosa Schaff.) which is reputed common in the eastern United States
and in Europe, S. laminosa Fr. of Europe, and S. spathulata Schw. (Ste-
reum spathulatum Schw.) (sparassis Herbstii Pk.) of America in the thin-
ness of its lobes and by its very pronounced perennial sclerotioid root-
stalk from which the sporophore develops annually (fig. 2). Since the
rootstalk is usually attached to the deeper lateral roots of its host, it is
often of a surprising length especially if a thick deposit of forest litter has
accumulated around the base of the tree. Specimens have been found
50 cm. in length but the average is from 20 to 31 cm. No record exists
of such a rootstalk for any other species. Sparassis crispa has a rooting
base but it is not known to be perennial. It is possible that this phase
of development is common to the other two species but has been over-
looked. Sometimes the underground stalk is divided into two secondary
ones each supporting a sporophore (fig. 2). The spongy character of the
upper portion of the rootstalk soon merges into a very hard, compact
mass and at the point of attachment to the root has very much the ap-
pearance of true cellular structure with the component filaments arranged
longitudinally. The periphery of the stalk at the surface of the ground
is composed of hyphae very much modified into a hard encrusting layer
and may sometimes have a resinous appearance. The mycelium at the
base of the stalk usually cements the earth into a hard stone-like body
often of large dimensions. The fungus has not been found growing in
the soil unattached to woody material. It is doubtful if it ever does so
occur. All specimens so far collected were found at the base of trees.
The structure of the rootstalk is not that of a true sclerotium although
it functions as such, is permanent and produces new sporophores from
year to year. The stubs of old sporophores are plainly evident on the
old root stalk (fig. 3) and as high as ten have been found on a single speci-
men. It was expected that the rootstalk would have great power of re-
generation. This was tested on July 3, 1915 by cutting off a half-grown
« Lloyd, C. G. Letter No. 44, note 51, 1913.
1917] Weir: Sparassis radicata 171
fruiting end, showing not only the evident polarity'' of the rootstalk but
that it is a reserve structure of considerable reproductive power.
The formation of sclerotioid bodies from which their fructifications are
developed is common to a number of Polypores. Chief among these
noted in western United States are Polyporus berkeleyi,^^ P. umbeUatuSf^^ P.
frondoBus^ and Lentinus sp., parasitic on the roots of conifers and probably
unnamed. The latter species has a true sclerotium." Lloyd^* lists the fol-
lowing species growing from sclerotia-like structures and separates them
as a distinct group of the section Ovinus of Polyporus: Polyporus tuber-
aster (Japan, China, and Europe), P. Goetzii (Africa), P. Sapurema (Bra-
zil), and P. Mylittce (AustraUa). Three other species also with sclerotia
but not included in this section are P. bdsilapidiodes (AustraUa), P. sacer
(Africa), and P. rhinocerotis (Malay).
The formation of sclerotioid bodies seems to be common to the Cla-
variacese. Some of the large species of Clavaria are observed to spring
from large globose masses which when sectioned exhibit a very compact
structure and are known to last over for more than one year. This has
been observed by the writer for Clavaria aurea, C. amethystina and C.
formosa. The members of the interesting genus Typhula always, so far
as observed, produce sclerotia from which the sporophore is produced.
In view of the fact that the sclerotia-forming habit seems to be more or
less common to the Clavariacese, together with the fleshy consistency of
the sporophores, flattened or cylindrical anastomosing branches, large
size of many species, amphigenous hymenium, constant in most genera,
irregular in others, it seems that this family is very well defined. The
removal of the genus Sparassis to the Thelephoraceae, which possesses few
or none of these characters, would be, it seems, an unnatural arrangement.
* Weir, James R. tFntersuchungen tiber die Gattung Coprinus. Flora n.s. 108:
301-305. 1911.
*• Weir, James R. Some observations on Polyporus berkeleyi. Phjrtopath. 3:
101-103, pi. 9. 1913.
Later observed by Lloyd, Letter No. 60, note 391, 1915; and Letter No. 59, note
306, 1915; and by Overholts, The Polyporace® of the Middle Western United States.
Washington University Studies 3: 23, pi. 2. 1915.
" Lloyd, C. G. Letter No. 58, note 277, 1915; and Overholts in The Polypora-
ceae of the Middle Western United States. Washington University Studies. 3: 24,
pi. 2. 1915.
" Fetch, T. The pseudo-sclerotia of Lentinus similis and L. infundibuliformis.
Ann. Roy. Bot. Gard. Peradeniya 6: 1-18, pi. 1. 1915.
" Lloyd, C. G. Synopsis of the Section Ovinus of Polyporus. 74-76. Oct. 1911.
Cincinnati, Ohio.
172 Phytopathology [Vol. 7
the disease caused by sparassis radicata
The observation that possibly some members of the genus Sparassis
are parasitic on the roots of forest trees has been made by others. In a
letter to the writer, dated January 7, 1916, Doctor Cotton writes: "Spa-
rams crispa has been found frequently, and from its intimate connectioo
with the roots of Pinus and other conifers we are strongly inclined to sus-
pect that it is parasitic/'
Kirchmayr/^ it appears, was the first to entertain the suspicion that
Sparassis had symbiotic or parasitic tendencies. Working with Spara$$iM
crispa, he found that the stalk of this species penetrated deep into the
earth at the base of the tree (Fohre). Boring into the roots from which
the fungus appeared to have sprung, he found that after passing througli
a sone of healthy wood the auger encountered diseased wood. TUs wood
was of a brown color, gave out a strong odor of turpentine, and was very
soft so that the auger readily pushed through it. Two trees when cut
showed that the brown rot extended up into the heartwood of the trunk
for a distance of two meters. The decayed wood could be nibbed into a
fine powder and gave out an odor of turpentine. The decay resembled
that produceii by Polyporus sidphureuSf the checks extending veitaeally
and |)aralleling the annual rings. The checks were lined wiUi a fine my-
celial layer which was encrusted with granules of calcium oxalate. Larfee
peicres of the cubical checkoil wockI could Ix^ removed from the hollow in
the heart wo(m1. The heart w(mx1 in the larger roots was also decajred,
while the sapwocxl was infiltrated with pitch {"verkieni'*). The decayed
wcHxl largely dissolved in anmionia prcxiucing a thick brown liquid whidi
on neutralisation held a brown deposit in suspension.
The author was unable to demonstrate the relation of the mycdium of
Sparannis crinpa with that in the disi^as^^l w(xxl. He calls attention to
the fact that the shrinkage of the wo<n1 in the fonn of cubes with surfaces
covennl with a fine white mycelial layer, bn>wn color, odor of turpentine,
and ability to Ix' niblMMl into a fine powder an* chanicteristic of the decay
pnnhi^Ml by PolyfHfrun HchweiniUii, In the writer's experience the rot
of l^olyfHtruA HchuTinitzii may not always Im» acrompanied by the pnxluc-
tion of s|)<>n>phon's until a long timo after the wcxhI Ls well advances! in
diTay. Sinre din^t ronn«M'tion of thr mycelium of the base of the sptiro-
phon' with that of the d(M*ay('<l winhI was not disc*emible, it seems quite
pn»b:ibl(* thut i\w invrstigator hii^ made an inc(»mH*t diagnosis.
A ran*ful examination bv the writer of six tn»es, the nx>ts of which lx»re
the fructifiration of SfHintssis nuUcntn has not n»vealiNl, with one excvfv
** Kirrhiiinyr. (Mmt «li*ti pHruNitiKiiiUH v<»ii Poly|>c>rui« frondoflus Kr. umi S|MirMi-
•u ranioiia .^luifT liodwifciii M: XW .'MT. 1914.
1917] Weir: Spakassis radicata 175
with a thick, mycelial mat from which the rootstalk takes its origin. The
greatest decay occurs at the point where the rootstalk is attached and
is at first confined principally to the sapwood. Small roots originating
from larger ones to which the rootstalk is attached are usually decayed
throughout. The early decay of the heartwood in the larger roots is
probably prevented by the large amount of pitch which they contain.
Eventually the heartwood is invaded but is pot broken down uniformly.
Elongated pits filled with a white mycelium are formed in different parts
of the wood, often anastomosing in such a manner as to leave long pieces
of partially decayed or solid wood which may be very readily removed
(fig. 5). Sunken areas on these pieces correspond to similar pits on
pieces which have become wholly disorganized. These elongated pits are
often bounded by a white mycelium arranged in the form of a network.
The tissue in the heartwood is brittle but can not be rubbed into a fine
powder as described by Kirchmayr for wood attacked by Sparassis crispa.
The rot of the heartwood is always of a darker color than that of the
sapwood. In Douglas fir it is brown; in spruce, of a more yellowish color.
Away from the seat of first infection the mycelium may advance into the
innermost heartwood causing the formation of a pitchy zone next to the
sapwood. The diseased wood may be drawn out of such roots in strips
leaving a hollow cylinder. The cambium and outer bast are always, how-
ever, permeated by the mycelium in the characteristic fan-shaped masses.
The action of the mycelium in the resin ducts of the bast causes a flow
of pitch which may cement the soil to the root in stone-like masses.
The fact that the fungus can maintain its activity in the cambium in
roots deep in compact, mineral soil is very unusual. Some of the root
fimgi which attack primarily the heartwood may follow the roots to a
considerable depth, and Annillaria mellea and Fomes annosus habitually
attack the cambium to a considerable distance in the mineral soil, but in
the experience of the writer no other species has developed this ability
to as great an extent as Sparassis radicaia. The decay is apparently
confined to the roots proper, never having been traced beyond the sur-
face of the soil. In case of an excessive accumulation of forest debris
around the base of the tree the decay may extend higher up on the lateral
roots than is ordinarily the case when this accumulation of materials
does not occur.
Only two species of fungi are definitely known to parasitise the roots
of coniferous trees in the temperate zone, viz, Fomes annosus and ArmiU
laria mellea, Rhizina inflata}^ may possibly be grouped here but in north-
western United States seems to be confined principally to seedlings.
^* Weir, James R. Observations on Rhizina inflata. Jour. Agr. Research 4:
93-05. 1915.
176 Phytopathology (Vol. 7
There arc a number of fungi which attack the roots of forest trees, are
not strongly parasitic and do not cause a rapid browning of the foliaiEe
and rapid death. Their action is confine<l mainly to the heartwood of
the roots and the base of the trunk. The most common of these is Poly-
porus schweinitzii. Other species which are either wholly confined to
the roots and bases of trees or extend into the roots from infection through
wounds on the trunk are Trametes Piniy Echinodoniium iindoriuni^ Poly-
parus sulphureuSy Porta trei'ni, and so forth. In the light of the pres-
ent status of the study it can not Ix? stated just how rapidly Spara^unn
radic€Ua causes the death of its host. It has not l)een found on repro<hir-
tion or young trees. The plant is not abundant but sufficient data have
been assemble<l to show that it may l)e placed in the same g^roup with
Armillaria mellea and Fomen annoaus.
To date only four trees, two Douglas firs (Pseiuiotsuga taxifolia), cme
white pine {Pinua monticoUi), and one spruce {Picea engelmanni) have
iKH'n found to have succuhiIkhI to the action of the fungus. The conclu-
sion that the <l(»ath of th(^sc» tn»es was causc*d by SjHira^Hts rndicala was
arrive<l at l)ecau.He of the absence of any other fungus or factor which has
heretofore l)een accre<lite<l as causing the death of trees. Several un-
healthy trees with the fungas on their roots have l)een studied, but the
common n>ot fungi wen^ prt^^nt making a cornH*t diagnosis impossible.
The fungus has in ev(»r\' ciis<», however, Uvn found to cause the death of
the living parts in the nK)ts to which it wjis attachcnl.
HOSTS AND DISTRIBUTION OF THE FUNGUS
Sparanms radicata is ver>' widely distributed in the Northwest, hav-
ing lH»on found by the writer in British Columbia, Washington, On^gon.
Idaho, and Montana. Sparasais crispa as report^nl from C^alifomia is
ver>' probably bascMi on this species.
The fungus has been found attacking the r(K)ts of the following conifers:
PnewUAifuga taxi folia, Picea engelnuinni, Pinujt moniicola^ and Ijorix (Kri-
dentalis. Its cK'currenc<» on the nK)ts of broa<l-leaf species has not U^en
note<i by the writer. Kirchmayr cites instances of the occurrence of
Sparannttis rrM/xj on oak an<l l)eech and other bn)ad-leaf species.
SUMMARY
The large s|)ecies of Sparassis in the wwtern Tnited States is found to
diffiT in a numlK*r of <ietails from Sptira^sis laminosa, »S. crispa, and N.
ttpathuUitn, and is clt^scrilMHl as new under the name Sparansis radicata.
The fungus is chiefly <iistinguish(Hl by its thin lol^es and an unusu-
1917] Weir: Sparassis radicata 177
ally large perennial rootstalk which is of the nature of a sclerotium and
from which new sporophores are developed from year to year.
The most important feature in the life history of the species is its para-
sitism on the roots of conifers. The myceUum attacks the bast of the
roots and later the wood, producing a yellow or brown, carbonizing rot.
Office of Investagations in Forest Pathology
Bureau of Plant Industry
Missoula, Montana
SOME CHANGES PRODUCED IN STRAWBERRY FRUITO
BY RHIZOPUS NIGRICANS
N K I L E. Stevens a \ d L o x A. H a w k i n a*
INTRODrCTION
Tho rot rauscMl hy Rhizopun nigricans EhrenI). in fttrawl)ern' fniitu
(Fragnriii sp.) and |K)tatc) tuJ)ors {Solanum (uberosum) has l)Ocn i«tu<iie<l
l)V the writers. In \Hy{\\ tlu»s<» cimk'h as w(»ll as in others' the rot caiu«eti bv
this funfai-*< in characteriziMl l>y the rapid softening of the affecte<l tiwuf«
aeeonipanied hy the loss of a Iarfi:e amount of juice. The morpholofQcal
chan^^s pro<hi(*ed in the straw!)err>' by Rhizopu.s nigricans have alreaily
Ix^en inv(*sti^ated. It was to (»Y>tain sonu* information on the bi<>-<*heini-
ral rhanf^'s brought al)out in the strawU^rry fruit by this funinu* that
tlie presc»nt study was und(»rtaken. In this work the effect of Hhizopu*
nigricans on the su^ar. a<*id, ix^ntosiin, and crude fat content and the
fXTct'iitage of dry matter of the strawU^rry was studied.
METHODS
The U»rries uwd in this study wert» all of the variety Missionary, itniwn
at Vienna, Virinnia. and picked lK»twe<»n May 2() and June 5, \9U\, The
methcNis for disinfecting an<l .*<amplinK us<'d suc(*essfully by one of the
writers in studit^ of fM'ach (3) and (M)tato (5) rots were found inapplicable
to this work. The outer layers of (vlls of the strawlierry were s<i injuretl
by antis<>ptics. such jis mercuric chlorid an<l alcohol, as to n»nder n*suUj>
of doubtful value. es|M*ci:illy as it had already Inhmi <ienionstnit4Mi that
under normal moistun* conditions the mynOium of the funfni^ |Cn»W!i
chicflv in the outer n'll la vers (S). The textun* of the Htrawl»errv. «f
coursi'. pn»vents |>ortions of the siiine U'rry U'inK u.*«<mI for incN'ulation
and r(»ntn»l. The* error dut* to variations in individual fruits inav lie
.«om<*what greater than where iMirtions of the same fniits can lie t^mifian'il.
' In th«* t\(MTiiiiriitf (If.NrriUMl in thifi paiMT the rultiirul work wan lionr by St«^
vi*nH. '\\\o jiiiiHir utithnr it* rf.«p<inHi)>lf* fi*r tli«* rhciniral work. Th^ wrtCem an>
inili-htcil t<» Mr A A. HilfV of iIh* uttwv nf DniK-IMant. Poiflonoun-Pltiiit. Phymolo.
Kiral. ami FiTnirntatinn InvrntifcationH for nHHiHtnnnf in the chemical work.
' 'I ht' litiTaturc r«>fi'rrinK to thr cfTcrt of lihiznpus uigricann <in various fruit«
haa \fvti l>ri4'tly rrvicwrtl in another pa|)cr tmon to U* piil>liiihf*d. Stcvena, Neil K.
and Wilrox. H. li . Uhixopuri rot of fitrawU'rri<'H in traniiit. l*. 8. D. A. Bui. 531.
1917] Stevens and Hawkins: Rhizopus nigricans 179
After some preliminary experiments the following method was found
satisfactory and was followed throughout the work: Berries as nearly
uniform in size as possible were picked when nearly ripe, i.e., when about
half of the berry showed a bright red color. They were picked early in
the morning while still cool and covered with dew. The calyxes were
removed and the fruit washed several times in sterilized, distilled water.
They were then placed in wide-mouthed flasks which had been plugged
with cotton and sterilized. Three or four berries were usually placed in
each flask. The berries were inoculated with spores and myceUum from
pure culture, a strain of Rhizopus nigricans isolated from strawberries
shipped from Florida during February, 1916, being used for inoculation.
The method usually followed in preparing the samples of fruit for analy-
sis was to grind the berries in a mortar and then wash the pulp quantita-
tively into the proper container. The flasks with the berries in them
were weighed immediately before the fruit was prepared for anal3rsis
and the washed and dried flasks were weighed again after'the berries and
juice had been removed. The wet weight of the fruit could thus be cal-
culated. All determinations were related to wet weight of the sound
or rotten fruit. The methods for the determination of the sugars, pen-
tosans, and dry matter were similar to those followed in the studies of
peach brown-rot (3) and the rots of potato (6).
The acid content was determined by grinding a sample of fruit, usually
about 20 grams, in a mortar, then allowing it to stand three days in a
flask with 150 cc. water to which a Uttle toluol had been added. The acid
was titrated with n/10 sodium hydroxid in this flask using litmus solution
as an indicator. The end-^point in these titrations was not as exact as
might be desired because the pigments of the strawberry which were pres-
ent in the solution made it impossible to detect slight changes in color.
However, the determinations are all comparative and the differences in
acid content in the sound and rotten berries are large. The crude fat
determinations were made by extracting the dried and ground samples
of fruit with water-free ether, which was then evaporated and the
residue dried and weighed.' A number of samples of freshly picked sound
berries were analyzed to obtain some idea of the variation in the con-
tent of the compounds determined between the individual samples. The
results of these analyses are shown in table 1.
From table 1 it may be seen that there is some variation in the content
of the compounds determined, especially the acids and sugars. The
' Wiley, H. W., ed. Official and provisional methods of analysis. Association
of Official Agricultural Chemists. As compiled by the conmiittee on revision of
methods. U. S. Dept. Agr., Bur. Chem. Bui. 107 (rev.), 272 p., 13 fig. 1908. Re-
printed, 1912.
180
Phytopathology
(Vol.7
TABLE 1
Showing the content of sugars, pentosans, acids, crude fats, and dry maUer infreski^
picked strawberries. Three separate samples used in the
determination of each substance
PBBCBIfTAOB
WCT WBIOBT
AOD AS CC.
KOBMAL ACID
rBB 100 OBAMB
WBT WBIQHT
RBDUCINO
SUOABII AB PBB-
CBNTAOB WBT
wxioirr
BrCROBB AS
PBBCBNTAOB
WCT WCIORT
CBUDC rATB AS
rCBCCNTAaB
WBT WCIOBT
OCT MATTSC
AC rscaurTACS
WCT WCMBT
0.57
0.59
0 56
2 59
2.10
2.15
2.89
3 67
4.03
1 37
1 69
1 65
0 27
0.29
0.34
7 97
8.17
8 15
variation in pentosan content is slight. There is a considerable amount
of sugar and the acid content is rather high. The l)errie8 of course weir
not ripe when picked and analyzed but wen* at al>out the stage of ma-
turity at which they are har\'e8ted in some sections of the South where
this variety is ^own commercially. A comparison of the results of the«e
determinations and the data obtained from the analysis of the sound
strawlH»rries which were analvzed three, 8(»ven, or fourteen da>'8 after
har\'esting gives some data on the (*hanges which take place in picked
strawl)erries during ripening.
In table 2 is shown the pentosan, acid, sugar, and crude fat content
and the percentage of total dr>' matter in the sound and rotted samplen
of fruit at different times after harvesting. The analyses are also of some
interest in showing the amount of the various sul>stances in the straw-
lK»rr>' fruit. (Considerable work hjis, of cours<». already been done on
this subject by various investigators. A review of much of this work
is given by Wehmer (9, p. 284-285). In table 2 the rc9ult« given are
averages of at least three determinations of the various compounds on
as many separate samples.
TABLE 2
Comparatire pentosan, arid, sugar, and crutie fat content and the amount of totai dry
matter in sound and rotten strauberries
rCWTOBAMB
4B rBKCRNT-
AOB WCT
WBKiirr
0 51 0 51
1
0 :js 0 .TJ
ATiD ro^rtBjrr
AH CC KOBMAL
ACID, rcB 100
oMfi. 09 rBrrr
li ' li
1 i :2
1 9H 1 H9
0 7r> 1 4'.»
0 73 1 .V>
BBOrclMQ
•lUABB
AB rBRCBNTAUB
WBT WB|(«HT
BrCBOBB
AS rBaCBMTAQB
WBT WBIUHT
CBCDC FAT
ooirrcirr
AC rcccnrrAOC
WCT WCtUMT
OCT MATTBC IC
•nLAWCKCCT
AC rccrciTTACS
W«T WCMVf
w
1 95 1 :Ci
0 44 ■ 0 13
li li
1- ' 1-
0 36
0 34
P
P
P
3
1
14
1 w» 0 fki
0 Jf) ' 0
0 40
0 31
7 19
««3
«19
5M
1917] Stevens and Hawkins: Rhizopus nigricans 181
From the results shown in table 2 it seems that the pentosan content
is no lower in the rotten fruit three days after inoculation than in the
corresponding sound samples. There is, however, a decrease in the
pentosans as calculated on a wet-weight basis after the fimgus has acted
fourteen days. It seems probable then that the fungus utiUzed a por-
tion of the pentosans. It is interesting to note that there is a somewhat
similar decrease in the percentage of dry weight in the inoculated berries
in two weeks so that if the pentosans were calculated on the basis of dry
weight at the time of analysis the percentage of the furfurol yielding sub-
stances in the sound and rotten fruit would be approximately the same.
The effect of the fungus upon the acids seems to be to reduce the acids
slightly, as the acid content after the first three days is somewhat lower
in the rotted samples than in the corresponding sound ones. The acid
content of the soimd berries decreased rapidly until at the end of seven
days it was only about half that of the rotted berries, and a similar ratio
k evident seven days later. From a comparison of the acid content of
the soimd berries in table 2 with that of the freshly picked berries shown
in table 1 it is evident that there is a gradual decrease in the acidity the
longer the berries are allowed to stand and that this decrease is much
more rapid in the first week. That this decrease was not due to a neutrali-
zation of the acid by ammonia either in the sound or rotten fruit was
shown by the negative results obtained from several series of ammonia
determinations by Folin's method. The acid is apparently used up by
the berry in its metaboUsm, probably in respiration. That the decrease
in acidity in the sound fruit was greater than that in the rotted berries
seems to indicate that the mechanism for the utilization of this acid
is destroyed or its action vihibited by the fungus. The fungus apparently
uses little of the acid.
The sugar content of the rotted berries is always lower than that of
the sound fruit of the same series of samples. A comparison of the per-
centage of sugars in the sound fruit, as given in tables 1 and 2, shows that
the sugar content rapidly decreases after the strawberry is harvested.
The sugars as well as the acids are apparently used by the strawberries
in respiration or in other metaboUc processes. The much more rapid
decrease in sugar content of the inoculated fruit is evidence that the fun-
gus uses the sugars.
The percentage of ether soluble material in the soimd and rotted ber-
ries, considered in the tables as crude fats, does not undergo any decided
decrease when the berry is rotted.
The percentage of dry matter in the rotted berries is less than in the
sound fruit.
As has been mentioned above, this strawberry rot is characterized by
182
Phytopathology
(Vol. 7
a rapid softening of the tissue, the loss of water, and apparently a itenend
collapse of the l^rry. It was considered of interest in this connectioD
to determine the amount of sugar and acid present in the juice which
escapes from the inoculated l)erries as compared to that in the juice of
healthy berries picked at the same time and maintained under the same
conditions of moisture and temperature. For this experiment the two
samples of I)errie8 were picked, washed in - sterilized water, and placed
in liter flasks. The l)erries in the one flask were inoculated with Rhiio-
pus in the usual way. while those* in the other flask were maintained is
controls. The flasks weiv filled to the wime height and were allowed to
stand in the lal)oratory for three days. The juice was then poured off
the inoculated fniit and the lx»rrics of the control sample were froipn
with carl)on dioxid, and the juicv expn^sstnl with a fniit press. The migar
and the acid content of the samples of juice from Iwth lots of berries were
determined according to the usual method. The results are shown in
table 3.
TABLE 3
Sugar and acid contrut of juice from nound and rotted utrawberrieM
nf-iiAK (rzmcr.sr\u%)
ATIDITT INTBMIi
i or iioBMAi. Mem
Juice frtMn kmiimI (niit Juice fruni ruCUtd fruit
Juice frtMn
•ound fruit
Jui»trom
rotlifti irmt
Rfvlucinc ■UfAT
SucnM* ' K«Hluctnc sucAr
Sucrraw
2.66
2 9S
0.62
0.14
0 191
0 208
DifTunion tension in Atmonphon^a.
jncB rmoM
•ocMD mcrr
0 604
8.37
iricB
1 037
12.50
From tabh* 3 it is appan^nt that the acidity of the juice from the di»-
c:i.s4m| lK*rri(*s is .nlightly higher than that of the juice from the sound oneti,
while th<* ."iugar content is considerably lower. In table 2 it is shown
that the sugar content of the dis<*a.^Mi fniit thnv days after inoculation
\va> <l<'cn'iL»*<M| consi(h»ral>ly Ih*Iow that of the control samples. This
may. of coursi', account for jmrt of the diffen»nce. Howex'er, the suirar
rimtent of till' ino<ulated InTries after thnn* clays is a little over half that
of the >4iun<i fruit, while the sugar content of the juice that leaks out of
tlie rotted fruit i> alMMit one-eighth that of the jui<*e expressed from s<iund
Urrie**. Th<* utilixation of th«' ^ugar by the fungus, then, can lianlly
.'irrniiiit eiitiri'Iy for tli«' lower sugar <*onteiit of the juic<* from the inucnilated
li«*rri«*> and much of the sugar must still n'lnain in the infected fruit.
1917] Stevens and Hawkins: Rhizopus nigricans 183
Other materials than sugars and acids are, of course, present in the
strawberry juice. In order to obtain some idea of the amount of substance
in this watery extract, freezing point determinations were made on the
two samples of juice. These determinations were made with a Beckmann
freezing point apparatus in the usual way. The depression of the freez-
ing point (A) of these juices below that of distilled water and the calculated
diffusion tension (7, p. 30-31) or osmotic pressure of which these juices
are capable are shown below:
The freezing point of the juice from the rotted berries is considerably
lower than that of the juice from the sound fruit. As calculated by this
method the solution from the rotted berries has, obviously, a higher dif-
fusion tension than that from the sound fruit. The juice from the rotted
berries then is a more concentrated solution of some substance or sub-
stances than is the juice from the sound fruit. From these experiments
with the juice from the sound and rotted fruit it is evident that the
juice which escapes from the rotted berries contains at least a part of the
soluble matter that is present in the cell sap of the berry before it is attacked
by the fungus.
DISCUSSION
The effect of the fungus upon the various constituents of the straw-
berry as shown in the foregoing pages is much the same as has been shown
for other fungi and other host plants in similar studies. Most fungi
apparently utilize the sugars in their hosts when growing parasitically.
This has been shown by one of the writers in the case of the brown-rot
disease of the peach (3) and some of the Fusarium rots of potatoes (6).
That the fungi sometimes lower the pentosan content of their host
when living parasitically has also been shown (4, 6). Rhizopus nigricans
apparently does not utilize the acids of the strawberry to any extent.
Some fungi are apparently able to use the acids in their host plant while
others are not, probably depending on the ability of the fungus to assimi-
late the specific organic acids that are present in the host. Behrens
(1, p. 700-706) has shown that the acids in apples can be used by fungi.
Sclerotinia, however, apparently had little effect on the acids in the
peach fruit (3).
In considering these results it should be remembered that at the time
of the first analysis, i.e., three days from the time of inoculation, leak
had progressed to an advanced stage. That is, the berries were flattened
and a large amount of juice had escaped. At this time, as shown by table
2, only relatively slight changes have taken place in the amounts of the
various constituents for which analysis was made. There is apparently
no difference in the pentosan content between the rotted berries and
184 Phytopathology (Vol. 7
sound berries of the same af^, and the difference in the amount of acid
present is very slight. Some reduction in the amount of sugar, liolh
sucrose and reducing sugar, has of course occurred but as the sugar was
probably chiefly contained in the cell sap this change offers no due a«
to the cause of leaking.
The bio-chemical studies have, then, served to confirm the conrlusioo
derived fruni the hu^tological study that the changes, detectable by the
methods followed, which have taken place in the cells of the strawljerry
at the time leak occurs are relatively slight. The histological study
showed that the cell walls of the 8trawl)erry are seldom pierced by thr
fungous hyphae and that the protoplasm of the cells is only slightly alterpd
in appearanc(\ the nuclei in particular retaining their normal appearaner
until the c<»lls are crushed.
In a<*counting for the losj< of juice which occurs in strawberries attacked
by Hhizopuji nigricann the only tenable hypothesis seems to Yye that thr
fungus so affects the protoplasm of the cells, perhaps by secreting soaie
toxin, that it is no longer capable of functioning as a semi-permeable
membrane. In this connection it is interesting to note that (lortener and
Blak<»sl<H» (2) have n»cently demonstrated the presence of a substance in
Khizopus which is extn»mc»ly toxic to rabbits. Whether the proti>pla«iD
of thi» strawl)erry is kille<i at once by the fungus or whether it is anr**
thetized an<l rendenni permeable to the material dissolve<l in the cell
sap is an op<»n (|uestion. Further investigations on this subject mrp
planned.
BiRKAr OF Pla.nt Indistry
Washimjton, I). (\
LITKHATrUK (UTKD
1. Hkhkk.ns. Joiiawkh. IVitrnf^o lur Konntnis dor (HMtfaulniii. CVntM. liakt .
AUi. 2. i: 7(«)-7(m. IKIW.
'J. (lOKTENKK. K. A., and Hlakkslkk. .\. F. (>lMu*rviitiontt on the toxin of Khiiop^M
nifcriranH. .\ni«>r. Jour. PhvKiol. 94: XA-lVu. 1014.
3. liAmxiNs. L. A. SoriK* rfTfrtH of th<* l»rown-rot fiinguii on the r(»mpo«ition of
thr jx'arh. Afiirr. Jour. liot. 2: 71 SI. HH.'i.
4. — Th<* utihiati<in of r«'rtain p<*iitoH4'H and roni|M>undN of pcntoum hy (tlo
H'lla rinfculata. Ainrr. Jour. Ii«»t. 2: :C'> .'iSS. 191.V
5. Thr dim*a»M* of |Mitat4N'H known an "Irak." Jour. \gr. Uciioarrh S: fU7
t\. -- I'ifTiM't of rrrtain K|M'<*ic^ of KuMariuni on tht* rompodition of thr p«»tAto
ixi\n'T. J«Mir. Aicr. Kf-MNinh 6: lH.VP.m. lUHi.
7. LiviN(«M<i\. H K Th«' nMf of ditTuHJon and o!«nioli(* pn*fuiurr in pUnta. 149 p
('hir.'iKo. hit>ho|craphy. p. '2. llNKi.
H Sti.\kn^. Nkii. K. I'athotoKical hiHto|o|cy of HtrawlM'rriotf affected by
of liotr>ti« and Khiz<ipui4. Jour. Agr. HriM^arrh 6: 361-306. 1916.
\t. \\i:iiyKit. <*\Ki. I)i«* Pflanz(*ni«tofTv .... \KVt p. Jrna. 1911.
WITCHES-BROOMS ON HICKORY TREES
F. C. Stbwabt
With One Figure in the Text
In Ontario county, New York, trees of the shell-bark hickory, Carya
ovaia, occasionally bear witches-brooms apparently caused by the fungus
Microatrama juglandis (Bereng.) Sacc. During winter, while the trees
are bare, the "brooms" are readily detected at a considerable distance.
They are typical witches-brooms consisting of compact clusters of short,
upright branches. They are of all sizes up to about two-thirds of a meter
in diameter in the bare state and, of course, considerably larger when
in foliage. As many as thirty "brooms" of various sizes have been ob-
served on a single large tree.
The leaves on the "brooms" are yellowish green above, and white and
mealy with Microstroma spores on the under surface. Usually, they are
smaller than normal and much curled. In mid-summer they blacken
on the margins, then wither and fall prematurely. The fallen leaves are
not replaced by new ones as happens with the cherry witches-brooms
caused by Exaascus cerasi. The branch bearing the "broom" is, usually,
considerably enlarged at the point of attachment of the "broom" and
often dead beyond the point of attachment.
The constant occurrence of Microstroma juglandis on the leaves leads
to the belief that this fungus is the cause of the "brooms." Almost every
leaf on every "broom" shows the fungus over its entire under surface
while the leaves on all other parts of the tree may be wholly free from
Microstroma. The presence of the fungus becomes evident as soon as
the leaves unfold in the spring. This condition of affairs is not rare. It
has been observed during seven consecutive seasons on a large number
of "brooms" on nine separate trees in three localities — Geneva, Canandai-
gua and Victor.
On the other hand, Microstroma jiiglandis has long been known as a
parasite on the leaves of walnut and hickory and is widely distributed
in Europe and America; yet its association with witches-brooms has not
been previously recorded. In fact, the writer has been imable to find any
published account of witches-brooms on hickory trees. The writer, him-
self, has occasionally observed M, juglandis on the leaves of hickory trees
which bore no witches-brooms. At Geneva, in 1916, this was of com-
[917] Stewart: Witches-Brooms in Carya 187
)f the leaflets were attacked by Microstroma. On an adjacent branch
-here was a larger "broom" every leaf of which was covered with Micro-
itroma, but none of the other leaves on the tree were affected. A later
examination, made on July 2, revealed no change in the "broom" except
hat the margins of the affected leaflets had begun to blacken. Further
>bservations were impossible owing to the accidental destruction of the
^oung "broom."
New York Agricultural Experiment Station
Geneva, New York
A NEW LEAF-SPOT DISEASE OF CHERRIES
Bert A. Rudolph
With Three Figures in the Text
In August , 1913, a conspicuous loaf-spot disease of sweet cherries was
ol)ser\'ed by the writer near San Jose, California, and specimens have
lM*en received from the principal cherr>' growing sections between Red-
lands, California, and Cor\'allis, Oregon. No mention of it has been
found in any of the literature examined. Descriptions were sent to F. C.
Stewart, Ceneva, New York; Howard S. Heed, BIacksbui|s, Virfinia;
M. B. Waite, Washington, I). C., and Donald Reddick, Ithaca» New
York, and all expn\sse<i doubt as to its occurrence in their localities. It
is lH»lieve<l to Ik; jx^culiar to the Pacific coast.
IVa<i, definite, circular spots from one to al)OUt 14 mm. in diameter
ap|K*ar on the leaves. The dea<l areas are a pronounced reddish brown
or chestnut to mahogany color, and are .sharply differentiated from the
living ti.Hsue, exten<ling from epidennis to epidermis and commonly marked
with a delicate, (*(>n(*entric zonation consisting of narrow lines and liarker
bands. St»vcTal spots may coalesce* to fonn one large one.
i )n the upper surfa<*e of the leaf may nearly alwa^'s l)e found a minute.
whitish gray pustule lo(*ated in the <*enter of the s|M>t, and about which
the zones an* concvntric. The pastules an' upliftcnl bits of epidermal leaf
ti.NSue, and a minute inM*ct of the family Chalcididae has regularly lieen
found in tliein.
Within the s|M>t then* may U» a definite, centnd. circular portion which
i> lighter than the n*main<ler. This inner an*a sometimes reaches 0 mm.
in diameter and is ochra<vus to f(*rnigineus (Saccanlo's Chromotaxia^
or it ni:iy U* entin*ly absent. The darker portion is ciLstaneiis to badius.
Th«* pw«tul«». if pn'H»iit. is Ineated in the eentiT of the inner, lighter area.
if •*ueh an nrvii nreurs. The nnd(*r sid(* of the s|M)t pn*sents a slightly
dilTtTtht <'n|nr frmn the up|MT. The inner lighter an*a is isaliellinus, antl
thi' retii.'iiiiiiiK |M»rtinii Irttfricius. iK'ing lighter than the com*s|H»ntiing
ana nn the up|MT surfaer.
Tlif «l«Md ti^^^ue n'niaiii*<i intact within tlu* leaf.
Soiiiftiiiir^ ihf s|M>t-« *ipn'ad niit in an irretrular, Mimewhat indefinite.
nn»*:iif-liki' fa*«hiori i tig. 1;.
1917] Rudolph: Leaf-Spot Disease of Cherry 191
The details of subsequent inoculations are recorded in the writer's
thesis, deposited in the library of the University of California. In all,
over two hundred inoculations were made on the following varieties of
cherries; Royal Ann, Richmond, Morello, Black Tartarian, seedling sweet
cherry and California Wild Cherry (Prumis ilicifolia Walp.). During
the winters of 1913 and 1914 experiments were confined to the leaves of
seedlings. In the spring of 1914, however, large numbers of inoculations
were made upon the various named varieties. The method of inocula-
tion was the same as that described earlier. Checks were made on one
side of the mid-rib and inoculations with a flamed needle on the opposite.
Not more than ten of all the inoculations made in this manner were nega-
tive. In most of the inoculations no bell-glasses were used, and the spots
developed quickly, although less rapidly than where covered. About
twenty other inoculations were made in lots of two to five by laying bits
of the fungus in agar on the leaves without puncturing or injuring them in
any manner. All these were negative. The greater part of this work
was done in a greenhouse without heat.
The fungus is particularly active when inoculated in the leaves of Cali-
fornia Wild Cherry. About three dozen inoculations were made on this
plant, and not more than five proved unsuccessful. Frequently arti-
ficially inoculated leaves were so badly affected as to be shed from the
tree. This was especiaUy conunon when the trees were kept under bell-
glasses. The spots produced differ greatly in color from those on the
leaves of sweet cherries. There is usually a circular, inner area in each
spot which is avellaneus on the upper side and isabelUnus on the lower.
The remaining or outer portion of the spot is isabelUnus, and the lower
side latericius. At times the spots may be a deep brown color, especially
when formed more slowly.
Microtome sections made of the freshly produced spots stained with
Congo red and methylene blue show best the action of the fungus. The
parasite is intercellular. The chloroplasts of the cells lying just beyond
the tips of the advancing mycelium are first affected and cannot be dis-
tinguished. The cells collapse and disintegrate rapidly as the fungous
threads come in contact with them.
All inoculations in the bark and wood of normal cherry twigs were nega-
tive. Over two dozen inoculations were made on stems up to 2.5 cm. in
diameter. The bark was first sponged with alcohol. Slant cuts were
made with a flamed scalpel and the infectious material placed beneath
the flap, or inoculations were made by puncturing the bark through the
spore-bearing material. Both types of inoculations were either left ex-
posed or wrapped with thoroughly boiled linen strips or bound in absor-
bent cotton. Checks were also made in the same way. The wounds
healed normally, the plants apparently being unaffected by the fungus.
102
Phytopathology
(Vol. 7
When inoculated in the leaves of other plants the results are often as
pronounced as in the cherr}' leaves. The leaves were first sponfced with
alcohol and the inoculations and checks were made in the usual man-
ner. The results obtained are shown in table 1.
TABLE 1
ResuiU of inoculations of leaves of various hosts with a species of Allernaria fi
cherry leaves
HOST
mtmvvn
OBMtBTAnolM
Apple (Pynis Malus L.)
Winesap
Newtown Pippin
Box Elder (Acer negundo L.)
Hungarian prune (Prunui4 domestica L.)
Wickson plum (Prunus triflora Roxb.),
(P. Simonii Carr.) hybrid
Orange (Citrus nobilis Ix>ur.)
King Mandarin
Loquat (Kriobotr>'a japonica Lindl.)...
Potato fSolanum tulierosum L.)
Avocado (Persea gratissima (iaertn.)..
Watermelon (Citrullus vulgaris
Schrad.)
Pearh (Prunus pc'rsica S. A Z.) Oaw-
fcird p<»ai'h
20
25
50
18
•6
20
21
30
10
40
All positive
All positive
All positive
All positive
All positive
Doubtful
Negative
Negative
All positive
All positive
Positive
Jjirge reddish spota
Large reddish «poU
Large brown spoU
Small brown spoU.
Developed slowly
Small brown spoU.
Developed slowly
Spots liarely Urfsr
than on therheeki
Spots no larger than
on the checks
Spots no larger than
on the checks
Large reddish brown
spotii
Black spots. Devel*
oped slowly
Irregular gray*
brown spots
In ^*m*nil it wiis found that the funj^is prcxluctMl its optimum ftrowth
in the If'uf tissue* when tlu* atniosphon* W!is moist and warm and sunliKbt
at a niiiiiriiuin. The men* shading of an infected leaf with a piece of
pajMT was found to fH'nnit the prcMJue'tion of largtT spots in a nhorter time
than whrn* the leavers were exi)os<»d to direct Kunlif2:ht. The fungus is
a typi(*al wound panu^ite. all incx'ulations on uninjured leaves ha\*iiig
failiMl. Whrn a young leaf w:is incK'ulated lH»fon» Innng fully developed
a shot-bolf I'fTtTt M)mrtim(v n'sulted on its exiuinsion.
The fungus gn»w vigorously on the conunon culture media, and its
more imfMirtant characteristirs an* as follows:
1917] Rudolph: Leaf^pot Disease op Cherry 193
On non-nutrient agar. Growth rapid, mostly confined to the surface
of the medium which is not discolored. A small amount of long, aerial,
dry, silky, gray-white myceUum is produced. Spores thinly scattered
over the surface of the medium.
On nutrient agar (containing meat extract, peptone and salt.) Growth
vigorous, a copious aerial, downy mycelium is developed consisting of
long, branched, silky, gray hyphae. Spores commonly produced in
greatest numbers in concentric zones which are dark green at first becom-
ing sooty black with age. The spores are oUvaceus under the microscope.
The agar is cleared of any cloudiness by the fungus as it develops.
On steamed rice. Growth vigorous. A snow-white, aerial, downy
myceUum first develops which darkens to a dirty, greenish gray with age.
A flesh-colored peUicle is produced upon the surface of the medium darken-
ing with ^e to black. The rice grains gradually become colored a Ught
yellow. Spores are produced close to the surface of the pellicle and be-
neath the longer aerial hyphae. They are pale oUvaceus imder the
microscope.
On bread and prune juice (Duggar's Fungous Diseases of Plants, p. 24).
Growth vigorous. A copious, downy, aerial mycelium is produced which
is dull white at first becoming a dirty greenish gray with age. Patches
of older parts of the aerial mycelium are often yellowish. A cream colored
peUicle is formed on the surface of the medium becoming black with age.
Spores develop close to the peUicle beneath the longer, aerial hyphae.
They are dark oUvaceus imder the microscope. The medium becomes
darker as the fungus develops upon it.
On steamed potato slants. Growth vigorous. A white, downy, aerial
myceUimi is first produced becoming a dirty greenish gray with age. A
peUicle is formed upon the surface of the slant and may be flesh colored
or greenish, becoming black with age. FinaUy the aerial myceUum usually
collapses, and only a black, shining peUicle is observed. After growth
has entirely ceased the plugs no longer react for starch with iodine but
give a good test for reducing sugar with FehUng's solution.
On beet agar. Growth vigorous. An aerial, downy myceUum is first
produced which is gray-white becoming greenish and finally black with
age. Cultures have a sooty, granular surface punctuated with whitish
hyphae in scanty tufts and occurring singly. The aerial myceUum may
or may not be somewhat zonate. Spores are inclined to be smaller and
decidedly darker than those found on other media, being oUvaceus to
fuUgineus imder the microscope.
On steamed cherry twigs. Growth vigorous. A copious, downy, aerial,
white myceUum is first produced which becomes a dirty greenish gray
to black with age, giving the cut siufaces a sooty appearance. The bark
194 Phytopathology [Vol. 7
ifl Hpareoly covered. Spores are proiluced in abundance close to the nit
surfacen of the twif^. The}' are somewhat smaller than thoee on vmriouf
other media and are dark olivaceus to fuligineus under the microscope.
On nat^el oranges (Sterilized by washing the surface with mercuric
chloride solution). When inoculate<l in moist chambers at the navel
end a black rot of the rag or pulp cells results which is identical with that
produced by Altemaria Citri Pierce (fig. 2). At the point of inoculatioii
an aerial mycelium develops which Ls pulvinate and gray-white at firat,
becoming a dirty greenish gray with age. The rind discolors becoming
olivaceus in a gradually increasing area around the fungus colony. Se\'eral
months after inoculation the aerial mycelium, having overrun the oranf^,
bleaches out and ultimately l)ecomes a l)eautiful pink. The whole fruit
gradually settles down with a soft, moLst rot.
On + 5 nutrient agar agar. This medium remains liquid due to the high
acid content. The i.solated colonies of ai'rial mycelium are whitbth at
first l)eroming s<M)ty black with age and rounded or hemispherical. On
titrating the incdiuin thn*e ww'ks after planting it wiis found to have lieen
re<luc€»d to -f 4. The color was changed from a light amlKT to a <leep
l)rown (fuligineas). This destruction of acid by the fungus was oliserved
on various other media. The average of four titrations was alwa>'s taken.
In genertil it was ol)s<Tved that tlu* color, shape and size of the spores
pnMluce<i on various me<lia may var>' slightly, but the most important
characteristics n*niain the same.
The fungim l)ears a striking similarity to Altemaria Citri Pierce and e\Tn
a closer n^lation-^hip to an Altemaria found on watermelon leaves. Purr
cultun»s of the thn»<» fungi wen* obtained by the single-spore methoii.
In drop rultun^s of 4-2 nutrient broth the fungi may \h* said to l>e ich^ntird
morphologically. Possibly the spon*s of A. Citri are slightly rougher than
those* of the other two, but this difTerenc(* was not found to \)o amslsmi.
The thn*<* fungi <'antiot Im* difTen*ntiate<i on nutri(*nt and non-nutrim
agar, and tlu* nits (inxhu'cd by them in navel orang(*s an* identical.
The cherry .Xltcniaria cannot Ih» distinguished from the wateniH*lo'
.Mterniiria on -f-'> nutrient agar, but is di.stinguished from A, (^htri c:
this medium. The latter jmMhK'es colonies which an* circular, whiti^
at tir>t. iNM-oiiiing gray with age. They an* also flat or depn*ss<H| wi
erater-like e<*nters which an* darker in color (gri.sc*us-oIiva«»us).
The cherry .Mternaria is distinguishtMi fn»m the oth€*r two Altenia
when inoculated on <*herrv leaves only by the size of the spots and t
rajudity uith wliitli they an* produ<*e<l. The watermelon Altenia
priM|im»s the >nialle>t >|M)ts. and they lievelop nion* slowly, but the tl
fen'MH' is very slight. Over four <lozeii ino<*ulations wen* made with
(Vri and the watermelon .Mternaria.
1917]
Rudolph: Leaf-Spot Disease op Cherry
195
The cherry Altemaria was distinguished from the other two by its ac-
tion on watermelon leaves (var. Cuban Queen), being the least virulent
of the three fungi. Sixteen inoculations each were made with the cherry
Altemaria and A . Ciiri on separate plants. Twenty-four inoculations were
made with the watermelon Altemaria on a third plant. The black spots
produced were identical, but the watermelon Altemaria spread to the stem,
killing the plant, while the cherry Altemaria confined itself to the leaves.
A. Citri formed sUghtly larger spots than the cherry Altemaria but also
confined itself to the leaves.
The cherry Altemaria may be further distinguished from the other
two in the matter of spore germination. Fresh spores of the cherry Alter-
FiG. 3. Sporophores and Spores of Alternaria Cithi var. Cerasi
From drop culture of 4- 2 beef broth with peptone. X 500
naria will germinate in neutral, +1, and +2 broth in less than twenty-
four hours, but in +7, and +8 broth germination rarely takes place in
less than a week. It will grow and produce spores in +8 nutrient broth.
Spores of the watermelon Alternaria will germinate in +8 broth and grow
feebly, but the fungus has rarely been observed to produce spores in this
medium. Spores of neither will germinate in +9 broth. A, Citri will
grow and produce spores in +6 broth, but spores will not germinate in
+7 broth. Spores of both the cherry and watermelon Alternarias ger-
minate in +7 and +8 broth with the greatest difficulty, the germ-tubes
developing in an abnormal manner and resemble the budding of yeast.
Ordinarily the germ tubes are straight with but few septa, but in broths
of high acidity, the opposite is the rule.
196 Phytopathology (Vol. 7
The cherry Altemaria cannot be distinii^ished from the others by
fcprmination testo in alkali solutions. All germinate readily in tap water
containing 8 per cent normal alkali (sodium hydroxid), but in —10 tap
water germination may not take place for a week. None of the three
fungi will germinate in — 11 tap water.
On steamed rice the watermelon Altemaria colors the rice ipiuns a
riightly lighter shade of yellow than does the cherry Altemaria, while
A. Citn colors them all shades of yellow to laterieious. In other respect*
the fungi arc identical on this medium.
On +5 nutrient agar the growth and acid destmction by the cherry
Altemaria was identical with that of the watermelon Altemaria. A
Citri within the same time only reduced the acid content 0.6 per cent
instea<l of 1 per cent.
Tlie longevity of the cherr>' Altemaria is largely dependent upon the
presence of moi.sture. However, sports ov(»r fourteen months old pro-
ducer! on steamed cherr\' twigs, which drie<l out soon after the planting
of the fungus on them, wen* found to l>e viable, although germination
rarely Vnyk [)laoe in less than a we<»k in neutml or +1 broth.
The various (»xix»riments enumerated show the three fungi to lie very
closelv n»Iated. The nanie of the watennelon Altemaria is not known,
but it is not lx*lieved to Ix* .4. cucurbitce Ia'X. which is also parasitic on
melon vines. The spon»s of A. ciicurbita' arc* descril)ed as longer and
narn)wer -mostly (>() to i\H by 8 to 9 /i — while the spores of this particular
Altemari^i an* only U) to 47 by 0.8 to 15/4.
The ch<»rrv .\ltemaria is not In'lieved to Ih» the same as Altemaria Certun
Potebniii found at Kharkov, Russia, the si>on's of that fungus l)eing dc^
cide<lly larger and pnMluced in v(4vety patclu*s on dr\' margins of leaves,
a(H*ording to Saccardo.
Believing this fungus to U' hitherto undesrrilMMl. and being of the opinion
that its rIoM' relationship to A. Citri Piera* entith's it to l)e elassificMi as
a variety of that s|M»('ies the writ(»r suggi*sts the name Altemaria (^itri
Pierce, variety Cvrasi with the following technical description.
Altemaria Citri Pierre var. Cerasi nov. var.
PrcN luring <lead >iMits on leaves of swe<'t cherr\'. Sjxits 2 to 14 mm.
in diameter, n'dtlish brown or chestnut above and lighter below, often
faintly zonat(% soni(*times with a distinct, lighter <*olore<l central area and
UMially ^ta^tinK from an uxm^vX injury, sometimes extending outward in
an indefinite nio?^ii<'.
\ wound pani>ite only, :ls shown by artificial infections, capable of
pnxluring s|M)ts on leaves of numerous plants. Distinguished from A,
Citri PiiTn* with difficult v.
1917] Rudolph: Leaf-Spot Disease of Cherry 197
Mycelium, in leaves of sweet cherries, sub-epidermal, of slender, sep-
tate, hyaline hyphae, 1 to 3 m in diameter. Aerial mycelium very rarely
produced and then only in the presence of unusual humidity and opti-
mum temperature, hjrphae 3.4 to 4.8 n wide, gray-white to pale oUvaceus,
long, silky, branched, septate; conidia, not observed in field, occasionally
produced in presence of unusual humidity and optimum temperature —
then close to the lower surface of the leaf, never on long, aerial hyphae,
clavate fusiform or elUptical at maturity, 16.3 to 57.8 by 6.8 to 15.3 /*, muri-
form, translucent, oUvaceus-brown, slightly verrucose or smooth, becoming
constricted at the septa with age, transverse septa commonly parallel,
3.3 to 9.9 fjL apart, with short, smooth, hyaline-subhyaline isthmus which
is usually 3.4 by 3.4 /x; several catenulate, in cultures 2 to 7 in simple or
branched series, somewhat variable in size, form and color, germination
from any cell, produced abundantly on various cultiu^ media. Conidio-
phores, short, in cultures 1.7 to 153 /* by 1.7 to 6.8 m, olivaceus-subhyaline,
erect, in general narrower than the vegetative hyphae.
Bureau of Plant Industry
Washington, D. C.
BLISTKU SPOT OF APPLES AND ITS RELATION TO A
DISEASE OF APPLE BARK
Dean H. R o b e^
With Three Ficjureh in the Text
In the present pufXT is descrilx'd a (ILseaHe of apples (fruits) which hit
\KH*n under investigation through two {^rowing seasons. A brief report
has already l)een published by the writer^ but so far no other mentioa df
it luis l)een found in the literature. There is fqven also a descriptaoil cf a
dis(*aseof apple bark which seems to l)e causally rt^ated to the fruit
The ex|)eriinental pr(K)f is not yet complete but considerable
that the relation actually exists has l>een obtained and will be found
marized in the s<»c()nd part of the papt»r.
BLISTKK SPOT OF APPLEH
Occurrence ntul general apiHtarance of the disease
The blister-s|)<>t dis<*:i.se was first noticed on July 6, 1915, on Noifoft
Pcauty (dwarf) as roughly (*ircular or sometimes irregularly lol)ed shallow
blisters, varying in color from light brown to black, and in siie from 1 lo
r> mm. in diameter (average aUmt 2 mm.) by 0.2 mm. in depth. Searrh
through the experiment station orchard at Mountain (irove, Miiwtmri.
thru .showed similar .s|K)ts on Melon, Ishewold, and Hawley. In 1916
s(>ots wen' first found aUxit the mid<lle of June on Blue Peamiain, Higfnn-
Iwitham, Yellow Transparent, Ii<'noni, Melon, Hock Pippin, I^Ansingburg.
Karly Hi|M*. Victuals and Drink. Isham, Ishewold, Hawley. Norfolk
Hcauty, Hr<l Astrachan, White Pippin (wrongly given as Yellow Xewt4»n
ui the rc|N>rt mentioned earlier).* Kloiidyke, Duling, ami Jonathan. AHow-
in^ for the fact that .su<*h varieties as Yellow Tran.^^pan'nt and Ke<l .VMra-
rliari wen' goiu' when afTe<*ted apples wen* di.scovere<l in 191o. this list
'Hhow> the diseasi' much mon* prevalent in the station orclianl in 19IH
tliaii m P.M.'). Little is known of its distribution. .\ppU»s showing typi-
' The writiT v%i>lirH to HrkiKiwIcd^c hi** iii«l('l>t4'<hirf(«4 to Mr. IlHTold Swartimt and
.\Iij»?» n«';itrir«' W liifc. uithoiit u ho*ti> <:in'ful :iii(l i*rtiri«*fit hflp thr work hrrr rr-
|»«irt«Mi roiiM rio* have iMM'ti :l^^oIl)pli^h4■<l m [\\v tunc HVailjihlr.
' Ho>M'. I)«';iii II llliHti-r >in»t of :ippl««'» ;i!i?*tr:irt . Phytopath. 6: II0 Krh
P»|ti.
200 Phytopathology [Vol. 7
appearance dcscrilxHl above (fig. i, B). On white or yellow apples such
as Yellow Transparent the spots are surrounded by a narrow greenish
ring; the same ring is found on Benoni and Duling, which are red when
ripe, but the spots on Red Astrachan show a red ring.
The disease is not a serious one in the way that bitter rot is serious.
Nevertheless, susceptible varieties often have the whole crop so seriously
blemished as to be unfit for packing in any but the lower commercial
grades. P'ortunately such severe injur>' seems to be confined to varieties
of minor commercial importance.
Cause of the disease
The disease is caused by an organism l)elonging to the genus Piieudo-
monas. It does not seem to have been descrilx^d previously and the
following name is propo8e<l and a description appended:
Pseudomonas papulans n. sp.
Morphological characteristicji. The organism Ls a rod, motile by one
to six polar flagella. Fiagolla occur at lx)th poles and were demonstrated
by means of Ixx'ffliT^s flngclla stain, using stains on Pseudomonas fiuart^
cens as check. (Px. fluorescena fumisheil by the American Museum of
Natunil History', New York). Staine<l from twenty-four-hours-old
agar cultun's the organism shows as a short ro<l with rounded ends, meas-
uring 0.9 to 2.3 M lofif^. by about 0.() ^i in diameter. It occiu« singly and
ver>' often in pairs. No si>ores or caj)sul(»s have lx»en \lemon8trat4*d. It
stains n*adily with carbol fuchsin, gentian violet, and methylene blue.
It is not acid fast and it den's not .stain bv (iram.
Cultural characterii<tia<. The following account is base<l on a study of
25 stniins of the organL^m isolatc<l from (1) naturally infectetl sp«»ls.
(2; artificial infections, and (3) infect ioas pnxluced with the n«iM>late<i
organism. Culture's wen* coinpanMl with li. coli and B. amyUnporus (fur-
nished I IV the .\iucrican Mummhii of Natural Hi.stor\', New York).
Thr organism does not form gas from |)eptonized Inmillon containinn
dcxtniM*. sn(M*haros4\ nialtos<'. hu'tose. glycerin, or maiHiit, and it d<M»!*
not rl<Mi<l the cl<KS(*d rnd of th<' fcrmrntation tuU* in any case. (tn>wth
sto|h« short in thr neck of the tulM*. indicating an obligate aerolM.\ The
oric:int>tii clouds lM*<*f bouillon -h H). slightly in twenty-four hours, and
nHMJiTatrly in forty-«'ight hours. It li(|U(*fics gelatin slowly at 20^1 \.
li(|uefa<*tion not U^ing complete* in test-tuiN* cultun^s until after twel\*e
to fourtn'ii w<H*ks. Some >t rains form a soft (*oagulum in plain milk.
S4»ine no roaguluni at all. but all of thiMu clear it in alnmt twenty ila\'ii.
The organism blues litmu> milk throughout <luring the first six day».
1917] Rose: Blister Spot of Apples 201
with the formation usually of a soft coagulum, and then gradually decolor-
izes it from above, with the production of a dark blue* color thr9Ugh-
out after sixteen to twenty days. On plain agar it produces a filiform,
slightly convex, whitish growth. On potato cylinders it produces, after
forty-eight hours, a whitish, filiform, irregularly spreading growth, which
after seven days shows a sUght browning, accompanied by a sUght darken-
ing of the mediimi. The optimuip temperature seems to lie between 25®
to 28*^C., though fairly good growth takes place at 20*^C. It does not
grow at 37°C. The thermal death-point has not been determined. On
dpxtrose agar and glycerin agar the organism produces a light green fluores-
cence as also in Uschinsky's and asparagin solutions, but it does not grow
in Cohn's solution. A test with six strains showed that the organism
grows in bouillon over chloroform and tolerates hydrochloric acid up to
+ 15 on Fuller's scale, and sodimn hydroxide to —5. The optimmn re-
action for growth seems to be about +10.
The organism is sensitive to sunUght. Petri dishes one-half covered
with black paper and exposed to sunlight, on August 3, on a bag of crushed
ice showed, for six strains, an average of 98 per cent killed after an exposure
of ten minutes. Six strains inoculated into test-tubes containing different
amounts of sodium chloride showed growth in all up to and including
4 per cent. Using the method described by Edson and Carpenter* it
was found that the organism, produces alkaU in plain milk during the first
ten days, then increasing amounts of acid up to forty days. On peptonized
bouillon containing two per cent of various sugars and alcohols it produces
acid from dextrose and saccharose, alkaU from lactose, and maltose, and
neither acid nor alkali from glycerin or mannit.
Quick tests for differential purposes; bluing of litmus milk followed by
decolorization from bottom upward accompanied by slow digestion and
the formation usually of a soft coagulum; fluorescence and luxuriant rugose
growth on glycerin agar; fluorescence on neutral gelatine but none on gela-
tine + 10.
Isolation of the organism. The organism was easily isolated from affected
apples by the method of poured agar plates. Spotfe were merely given
a good washing with steriUzed, distilled water, sometimes preceded by a
brief rubbing with a finger dipped in alcohol. The diseased material was
then scraped off with a sterilized scalpel and dropped directly into melted
agar. The colonies appeared in from thirty-six to forty-eight hours,
usually in pure culture. They were thin, smooth, circular, glistening,
• Saccardo, P. A. Chromotaxia seu Nomenclator Colorum. 1-22. 2 pi. Pata
vii, 1894.
* Edson, H. A. and Carpenter, C. VV. Micro-organisms of maple sap. Vermont
Agr. Exp. Sta. Bui. 167: 321-610. 1912.
2()2 Phytopathology [Vol. 7
whitish by n^flerted liRht, bluish by transinitted liRht, and 0.1 to l.U
mm. in diameter. Not all the affected varieties were used in thia work
but no difficulty was experienced in obtaining pure cultures from thoete
that were usetl.
Inoculation, l^in^ sul>-cultures from single colonies, the disease hah
l>een repro<luced on six varieties of apples. In this work 123 apples were
use<l, 60 of which were che<*ks. All of them were treated cm the trees
and were bagfced after treatment. The incubation period averaised about
fourteen days, thouf^h some apples failed to show signs of the disease
until the end of eif^hteen to twenty-five days. Some strains of the orfcanisin
were infectious on all the varieti(»s teste<l, others on only one or two.
Further work is nc<*essar>' to clear up this situation.
Inoculations wen* made in thret^ different ways, using twenty-four to
fort y-t»ight -hours-old iKuiillon cultun»s:
1. By spniying uninjunnl apples with th<' bouillon culture. No info^
tion n*sulted.
2. By spraying apples whi(*li had first Innm pricked with a flamed needle.
Seventy-two |kt cent infection resulted on Yellow Transparent, Jonmthan,
Melon, Hawley. and White Pippin. No signs of infection appeared on
I^*noni.
3. By hy|KMlennic inje<*tion just under the epidermis. Eighty per
e<>nt infection resulted on Benoni, Hawley, Jonathan, Melon, and White
Pippin.
Ino(*ulations with n*isolat4Ml (uiltun's by hyixKiermic injection on ^liite
I^ppin, Jonathan, and Melon wen> also successful. The organism isolated
from infections agn>ed in all characteristic's, morphological and eultural,
with the one us(mI for ino<*ulation. Checks to correspond with the three
methods of inoiHilation descrilM'd al>ove nnnaineii healthy throughout
tlu* s(*ason.
KorcSII-HARK OK SC'CKFY-HAKK (WNKEK OP APPLBS
DeAcriptitm of the f{ij<ease
What might U* called tlu* ({uiescent st:tge of the scurfy-lmrk disease
<M*rurs :us patches of n>ughen<Ml scaly bark which simiewhat respmhie
blotch ranker {PhyUostictti suUtnrin K. and Kj. but differ fmm it in show-
ing no blotrh pycnidia and usually no regularity of cracking up and down
or arri»NS X\\v limb (fig. 2. .1). Thes<* roughened |):itch(*s var>' gn*atly m
M/e fn»ni tliosi' covering only a few sipiare centimeters to thost* covering
till* wliolf Hjdt' nf a limb for a meter or in<»re. Thev an* usuallv f(»und i»n
thf iiurth ^ide of a limb and with few exceptions an* bordenNi by a pimpltMl
1917] Rose: Blister Spot of Apples 205
scaling off of small patches without the formation of a definite spongy
layer. Sometimes the diy, brown vestiges of such a layer can be foimd,
sometimes not. Possibly in such cases it develops slowly and progres-
sively from one point to another, loosening the bark only a little at a
time and drying down almost immediately.
No detailed studies of the diseased tissues have been made. A pre-
liminary survey of both free-hand and microtome sections shows the pres-
ence of peculiar spots or pustules like those described by Hewitt* in his
work with a new disease of apple bark in Arkansas and by the writer*
in work with what is almost certainly the same disease, known at this
Station for over ten years under the name of pimple canker. It remains
to be proved, however, whether this disease is a phase of or in any way
lielated to the true scurfy bark canker. The most that can be said at
fvesent is that they show considerable morphological similarity and are
often though not always associated on the same tree.
Besides the deep-peeling type of scurfy-bark canker there sometimes
occurs a "shallow-peeling*' type, in which only the epidermis is loosened.
A spongy layer is formed here also, but it is thinner and more evanescent
tiian in the deep-peeling type.
Occurrence of the disease
The disease occure on such standard varieties as Ben Davis, Jonathan,
Logan, White Winter Pearmain, Beach, Stayman W'inesap, Munson,
and Marsh, but much more severely on certain dw^arf varieties.
The writer has found the disease in numerous orchards in southern
Missouri and has received specimens of it from perhaps a dozen localities
scattered over the state. No information is at hand as to is occurrence
in other states, unless the trouble investigated by Hewitt in Arkansas be
considered a phase of it. Affected trees are not quickly killed as in the
case of Illinois canker — caused by Numviularia discreta — but there is no
doubt that the peeling off of fresh layers of bark every spring is definitely
injurious to the tree, aside from the opportunity given for entrance of
canker fungi and various bark insects.
Cause of the disease
The disease seems to be caused by the same organism as that of the
fruit spot. Poured agar plates, using material from (1) the deep lying
* Hewitt, J. Lee. An unknown apple disease. Arkansas Exp. Sta. Bui. 122:
481-491. 1912.
• Rose, Dean H. Report of the Pathologist. Missouri State Fruit Exp. Sta.
Rept. 1913-14 (Bui. 24): 30. 1914.
206 Phytopathology f\*OL. 7
spoDfor layer, (2) the pimply ridge at the edge of roughened areas, (3)
areas exposed naturally the preceding season when the loosened layer
peeled off, (4) the spongy layer under loosened epidermis, gave pra<rtically
'pure cultures of an organism very similar, morphologically, to the blister-
spot organism. Extensive cultural studies show, however, that instead
of one organism there are two different ones or possibly two closely related
but distinct varieties. One of them, represented by five strains, shows
great similarity in morphological and cultural characteristics to the bliiH
ter-spot organism; the other, represented by fifteen strains, resembles
the blister-spot organism morphologically but differs from it in cultural
characteristics. It liquefies gelatin rather rapidly, it produces a green
fluorescence on nutrient agar and it begins to clear litmus milk inside of
twenty-four to thirty-six hours. The first and second of these are not
characteristic of the blLster-spot organism, while the last is characteristic
only of three strains.
Inoculations
Inoculation of bark with the rapidly liquefying bark organism produced
swollen spots 1 to 2 mm. high and covering an area of roughly 1 sq. cm.
(fig. 3, C). At these swellings the typical signs of the disease were re-
produced, in miniature, and an organism was recovered which agreed
in cultural and mor|)hol(>giral characteristics with the one use<l for inocula-
tion. The check.s showed only a slight swelling. Typical blister spots
on the fruit wen* pnxiuced by inoculating Jonathan and Melon apples
with l)oth bark organisms and these in turn were recovere<l fr«>m the
lesions pr<Hluce<l. While such evidence is strongly suggestive it is not
final pnK)f, an<l nion* work is nec<\ssar>' l)ef()re the true relation U»tween
th« fruit di.s<»jLS<» and the bark di.*^»jis<» can Ih» dis«rovenHl.
In table 1 arc? shown the results of compamtive tests made with the
blLster-s|>ot organism and the rapidly li(|U(*fying bark organism.
A similar comi)arison lM*tw<H*n the bark and* the blister-spot organL^n
on the onr hand an<l pHcudomorum fluorescent on the other .show s<i many
<liffrrrnei»s that the two fonner must Ik» considrn»d entin»lv distinct fn>m
the latter.
Further work on these <iLs<':i.ses should include (Da continuation of
cultunil studi(*s of the blister-s|H)t organism and the two bark organbuiut.
(2; (To>s-incMnihitions from bark to fniit and from fniit to l>ark, uii a
study of the time and nxxle of infection and (4) a study of the microscopi-
cal eli:iraeteri>tirs of health v and <lis<»as«Ml tissues.
I
1917]
Rose: Blister Spot op Apples
207
TABLE 1
Comparison of the blister-spot organism and the bark organism
Agar plates. .
Agar slant. . .
+ 10 bouillon
Gelatin stab .
Neutral gelatin stab.,
Litmus milk
Sterile milk.
Glycerin agar.
Uschinsky's solution.
Nitrate reduction
Indol test
Ammonia test
Resistance to sunlight.
Flagella.
BLr8TKB-«POT OBOAKISM*
Whitish colonies, slightly
bluish in transmitted
light ; medium not
greened
Growth rather slow, no
fluorescence
Clouding and pellicle; one
strain showed faint green
fluorescence in 2 weeks
Liquefaction slow, begins
in 2-3 days. Complete
in 12-14 weeks. No fluo-
rescence
Liquefaction slow. Slight
fluorescence
Alkaline reaction for 10
days, then slow increase
in acidity up to 40 days.
Medium cleared in 2-3
weeks; final color blue
throughout. Banded
appearance produced by
3 strains
Clearing begins in from 7-
10 days, complete in 17-
20 days
Growth vigorous, elevated,
contoured to rugose.
Moderate fluorescence
Clouding moderate; bluish
green fluorescence. Pel-
licle of pseudo-zoogleae-
like fragments
None
No indol in 20 days
Ammonia produced
Average of 98 per cent
killed after 10 minutes
exposure
Bipolar; one to several
BABK OBOAmSIC*
Whitish colonies, slightly
bluish in transmitted
light; medium slightly
greened
Growth rather slow, fluo-
rescence marked
Clouding; green fluores-
cence after 1 week; four
strains showed pellicle
Liquefaction more rapid,
begins in 24 hours, com-
plete in 2 months. Slight
fluorescence after 1 week
Liquefaction more rapid.
Marked fluorescence
Slight alkaline reaction at
top in 2 days, acid reaction
below. Acidity increases
as digestion proceeds.
Banded appearance re-
sulting in clearing in 3-4
weeks. Final color green-
ish blue above, buff below
Clearing begins in 36 hours,
complete in 3-4 weeks.
Green fluorescence
Growth vigorous, elevated,
contoured to rugose.
Fluorescence more
marked than for blister-
spot organism
Same as for blister-spot
organism except fluores-
cence more marked with
some strains
None
Indol present in 10 days
More ammonia produced
than by the blister-spot
organism
Average of 50 per cent
killed after 10 minutes
exposure
Bipolar; one to several
^ Under this heading are included also the slow liqueflers from bark.
* Under this heading are included only the rapid liqueflers from bark.
208 Phytopathology (Vol. 7
METHODS AND MKDIA
As standanls for methods liiid for the making of media the writer has
followed directions given in Krwin V. Smith's Raeteria in Relation to
Plant Diseases, Vol. I, and ICyre's Ha<*teriolo|dnil Teehnie, 2nd Kd..
except where noted otherwise in the t(»xt.
Sl'MMAKY
1. In tlie foregoing i)a|K»r is d(\scrilHMl a l)aeterial disease of applf>.
no mention of w)ii<*h has Ihm'Ii foim<i in phytopathological literatunv
2. By isohition, niltural, and inoculation work, it is proved that thbi
dis<»ase is <'aused l)V a motile organism, which H(jueties gelatin slowly,
and lK»longs to the grccn-fiuorescent group of l)acteria.
'A. H<*caus4' of the Mister spots produc'(Ml by this orgsmism on the Mir-
fa<*e of apples, the naiiH* Pscmlofunntis pdjnthins is profKwed.
•I. Description is also given of a rough-hark or scurfy hark canki-r
from which has Im^cii isolated an organism also U'longing to the gn-«*n-
fluoH'Scent grou)).
.">. Kvidence is present I'd that then* are really two varieties of the hark
organism, one of whirh has all of the cultural rharact eristics of the hlister-
sfxit organism, including s|(»w Ii(iuefa<*tion of gelatin, while the c»ther dif-
fers fn»m it in S4'veral import .iiit particulars, including rather nipid li«|Uf>-
facti(»n of gelatin.
(>. Inoculation of healthy apple hark with two strains of the nipi<ily
lifiuefying hark organism pHxluccd small lesions which showe<i the typi-
cal <'racking Ioom* of diM*:iM'> from healthy hark, and in sevend e:is*'s thf
lumpy apjM'arance characteri>ti<* of the early stages of the scurfy -tiark
canker.
7. .\n organism agrei-ing in cultural react ion> with the one umhI fur
inoculati<»n has Imi'Ii re<*overed from thesi* le>ions.
S. Typical hli>ter s|)ot> have Ihh'Ii produced hv inoculation of Imth
tyjM's of hark organism> into healthy apples. From th<^» s|)ot,s the or-
gani>ms u>ed for inoculation havi* U-eii recov<»nMi.
\K \ preliminary <Mimparative >tudy of the cultural chaniet<Tist ii-^ oi
the Mi«»ter->pnt organi*»m ami the two h.ark organisms suggi^ts tluit fhf
difference iM-twei-n them are difTeren<-e> of degn-e nit her than of kind
That i^. that .all thre<* are iM»«»sihly men-ly varieties of one s|n»cic»s. Mon*
work i-* nec4'*.sary. however. U^fore this <|ue^tion ni n lati<inship can U*
M'ttlrd.
Mis-^oi HI Statk KiiMr l*!xPKHiMK\r Sr\rio\
M«>l\rAlN <IUoVK. Ml^S4U KI
THE PATHOGENIC ACTION OF RHIZOCTONIA ON
POTATO
H. T. GOssow
With One Figure in the Text
It appears that satisfactory evidence has been lacking clearly demon-
strating the pathogenic action of Rhizoctonia (Cortidum vagum B. & C.)
on various host plants, particularly the potato.
The attention which the well known disease has received in the new
world and more recently in the old, left, in my opinion and in that of
quite a number of other investigators, several important points unex-
plained. Every pathologist fully recognized the symptoms of this dis-
ease which have come to be regarded as typical Rhizoctonia infection.
The unmistakable folding of the leaves of the growing plant, together
with the brown stem lesions so frequentlj*^ described, in other instances,
or in addition perhaps, the formation of aerial tubers and peculiar small-
ness of subterranean tubers, are now well known as general symptoms
resulting from an attack of Rhizoctonia.
Indeed, the folding of the leaves associated with this disease differs
greatly from the curling of the leaves of plants affected with leaf roll,
particularly noticeable is this difference when examining the lower leaves
of an affected plant. Where, however, doubt existed, the presence of
stem lesions was looked upon as final proof of a Rhizoctonia infection.
I must confess, however, that the often surprising scarcity and appar-
ent superficiality of these lesions, nay, often enough their entire absence
in what was otherwise unmistakably a plant infected with Rhizoctonia
and not with leaf roll, frequently caused me surprise and certainly failed
to readily convince the farmers on the occasion of field demonstrations
of the correctness of the diagnosis — which at times I shared much against
my own belief.
For some time I have endeavored to discover the true pathogenic action
of this fungus on the host plant. My colleague, Drayton,* demonstrated
to my satisfaction the profuse permeation of the tissues of and surround-
ing the lesions, but even that failed to convince me entirely of the cause
of the characteristic symptoms, which statement is not intended to infer
^Drayton, F. L. The Rhizoctonia lesions on potato stems. Phytopath. 6: 59.
1915.
210 Phytopathology [Vol. 7
that I (louhtcHl Rhizoctonia to l^e associated with the same. But from
what evidence was available, the actual injuries caused by the stem le-
sions ^-ere so infinitesimal that it was felt the true injury is done elsewhere,
and what we did observe was the result of such unlocate<l but far more
serious injury.
A careful stud}' of diseased plants in the field revealed at first little or
no additional clues. Lesions were sometimes present where the lea\*es
were folded, the tubers were covered with more or less numerous lumpy
funfcous nuisses, indeed the roots often showed the well known pseudo-
sclerotia. Pot experiments showed the presence of sclerotia on rootlets
more abundantly th^m was the case in the fiel<l, and yet while abundiuit
superficial and lesser amount^s of intracellular h>^hae of Rhizoctonia were
found on microscopical examination, the evidence of an all round general
soundness of the underground parts examined still left the seat of the
injury undetermine<l.
On careful examination of the root system of a plant clearly affected
with Rhizoi*tonia and no other disease, that had lx*en pulled up from loose
sandy soil, or had l)een lifted with care by means of a fork or spa<le. one
factor at last attracted my att<»ntion, which later led to interestin^c oliser-
vations. This was the almost entire al)sence of the fine fibrous rootlets.
so common in sound plants. Surc»ly such rootlets must have been presu
ent ori^nally? When examininfc thereafter plants in various stagi*s of
infection one could obstTve a corresponding absence or presence of finer
nH)tlets ucconiing to the amount of disease present. Of course, in thi*
detennination can* is net»essiir>', but after some experience one cannot but
recogniz<» the existing n^lation of nK)tlets to deja"ee of disease.
What if the fungus a<*ted upon the rootsS of the growing plant similar
to the way in which it (1<h's on the r(M)ts of Rhizoctonia-infecte<l tulM^m
sprouted in a closed sten<ier <lish? The disastrous effects of the fungu'^
at the early stages of growth arc sometimes so pronounce<l as to kill off
growth liltogether; this is a H*ell known fact.
Ia'X us iM'ar in mind that in a potato field we find many stages of severity
of Hhiz(M*tonia infection, from total **mi.ssc»s'* to one, two or more shoots
clearly afIe<-t<Ml uj) to the cas4» wlu^n* the plant Invars plenty of aerial tulier*
and nuMHTous littU* |M)tat(M\s underground fnnn which the popular namr
**littlr jMitato (lis<»ji.*<<»" has sprung. .Verial tul)ers have l)een commonly
ji.H>ori:iteii with Hhizoiioiii:!. tlu'V are jxTluqw exclu-sively manufacture«l
from in.itrrial produteil l»y the heaves, a <*oinpanitively slow proresw, but
alua>> iii(ii(*:itiim iiiipainMl nnit function in plants where they may br
c<ni>i«h'nM| abiioriiial. .Verial tuU^rs natunillv mav occur from anv
• • •
raii.M* (MittiiiK otT or intrrnipting nN>t function. Imt only when such inter-
nipti(>n> an* gradual. We have, then'forr. no aerial tulN*rs in the black
1917] Gtjssow: Rhizoctonia on Potato 213
but the roots infected with sclerotia remain in the ground, since they are
not pulled up by the digger or are at any rate returned to the ground. With
the diminishing food supply in these roots, sclerotia develop ready for
subsequent attacks. This observation also accounts largely for the soil
contamination and the persistence of the organism iQ land once infected.
It also indirectly suggests a new means of control, viz., the prevention of
infection by cultural methods or the application of fertiUzers producing
vigorous plants in the first instance and aiding in the production of a
generous supply of new feeding roots.
Central Experimental Farms
Ottawa, Canada
212 Phytopathology [Vol. 7
leg disease, whore the cutting oflF of supplien is rather sudden. .\II th
symptoms are the lof^ieal results of the al)sence of the abundant fee<linc
n>ots. Roots are present in all growing plants, otherwise the planbt
would have died ; small and fine roots are less in evidence in affecte<I plant.*<
while a generous supply exists in healthy strong plants.
De<iuctions — however logical they may l)e — still are h\iM)theses and
hypotheses are not facts, but the accompanying plate will pmvi<ie
some foundation for the observations n'corded and may stimulate
wider n»s<»arches on this point than hiive l)een made so far. I am satu*-
fied fnnn the obs<»r\'ations made, that the destniction — often ver>' gradual
— but very jx^rsistent all the same, of all or many of the feeding rootj* of
the potato plant a<*c()unts for every one of the symptoms associatcNl with
this dis<»as4». The l(»sioiis which have so often lH»<'n recorde<i are evident I v
not of MTious conscHiuence, as indicated by their general superficiality and
fn»(|uent (»ntire al)s<»ncc. In some instan<Ts iiidcHMl these k^icms an* not
due U> Hhizortonia at all, but to Aciinomya's scabiej< (liissow, whirh I
hoiM» to sliow in anothrr pa|M»r, when they afford easy resting pl:i(*es in
the unprotecte<l su|HTficial cells for the mycelial m:iss4^ of I{hiz<N*tonia
sh(»wn in .Mr. Drayton's photo-mi<Tographs, :is well as for the [)c*rnie:ition
of the hyphae into the interior, which, as must have Imh^ii noticvd. is not
accompanied by any prominent injurious action ufMm the ctIIs invadtnl.
A stu<ly of Mr. Drayton's slidf»s clearly confirms this observaticm :is well
as the photographs ma<l<' fn»m them which an* ac<rssible to our readers.
The i)athogeni(* a<'ti<»n is as follows: We an* aware of the very pntfu****
growth (»f mycelium of Hhizo(*tonia. particularly in the dark, :u< al>«» of
the pHNluction of enormous <{uantities of ps^'udo-scltTotia on nMit> and
tuU'rs. Whethrr thr sclcrotia an* lt»ft over in the soil from pnM*edinK
(Mitato cn)ps or c»ther host plants, or whrther they have Uvn intnMiu<*«*«i
by untn^attMl infc(*t4Ml s4»<mI potat(M*s — (and what **farmers' nm'* [M»tatm*s
an* not infected?) <I(m»s not matter much. Th«» ti|)s of the fn-sh nnitlft^
niHiU fall a victim to tlu* invading myc<'lium. the nN>t cap iH'ing undoubt«Hlly
the nio*«t vulnerable |M»iiit and s<»on the short nM)ts have lN*<*n d«*>triiye4i.
the niyeelium meanwhile reaehes older rootlets, whieh it mueh niorv
ran*lv de>tro\>. though that hiis occurn'd. l)ut when* the mvcelium fn-
• • •
(luently prtHJueo re^^ting niyrelial mas.^<^s from which invading hyphae
iv^ue aInio>t >iniullaiieously with new nnitlets whieh an* pnMlui*«t| bv
the pl.mt in i\^ elTort to ree«*tab!i>^h its resources. This pnNH^s got-^ i»n
fc!r:idually and >lowly or more ra))idly de|x*nding naturally \i\Mm the vigor
of the plant. I'iiially the |NT*«i>tent efforts of the fungtis n*sult in deen-ik'^
uig Mf-NN. Ml ftii'^tratinL; the growth of the t ulnars. lN>cau.s<* of lack of Um^i
su|>phe'^ from the roots. :nu\ eventually in the production of m*rial tuliers.
.Meanulule harve>t-time ha> arrive<l, what tuln-rs an» there, are har\*este«l.
1917] G^ssow: Rhizoctonia on Potato 213
but the roots infected with sclerotia remain in the ground, since they are
not pulled up by the digger or are at any rate returned to the ground. With
the diminishing food supply in these roots, sclerotia develop ready for
subsequent attacks. This observation also accounts largely for the soil
contamination and the persistence of the organism iQ land once infected.
It also indirectly suggests a new means of control, viz., the prevention of
infection by cultural methods or the appUcation of fertilizers producing
vigorous plants in the first instance and aiding in the production of a
generous supply of new feeding roots.
Central Experimental Farms
Ottawa, Canada
SYNTHETIC' C ULTURE MEDIA FOR WOOD-DESTROYINT.
FUNC.r
Ernbst J. PiEPER, C. J. Humphrey and S. F. Acrbb
Wood-destroying funfd grow readily on many of the ordinarj' culture
media which have as their principal ingredients malt extract or plant
decoctions and meat extracts, usually hardened with agar-agar or gelatin.
As an example of such media, a malt extract preparation of the following
formula has proved very satisfactory for gt»neral culture work:
Extract of 1 pound lean }k>cP in distilled water 1000 cc.
Malt extract 25 grmmfl
Agar-agar 20 grama
This hiis found a wide us*» in Europe and luis \)een emplo^Td for much
of the routine work in the piithological section of the Forest PhmIucUi
I>iilK>rator>'. In certain lines of investigation, however, such as the test-
ing of the toxicity of chemical sul)st.iinc<»s, and in compiirative Xe»Xs on
the physiological U'havior of w(M>d-<Ic^troying fungi, this medium ht^
the s<Tious disa< I vantage's of lx*ing chemically complex and variable.
Its composition and constitution <lepend upon the nature of the meat
and malt extract, the method of preparation and the duration of stan<iing.
and physically it may 1k» variably <'oIloidal. Tlie |M)ssibility of chemi-
cal or physical combination of c<»rtain pr(»serv'atives with the highly com-
plex organic cotn|M>unds and also the coagidation of the latter by elec-
trolytes an* of cxtn»me importance also in toxicity work.
In the pn*s(*nt work an attempt was made to prt^pan^^ a synthetic mcHiium
which wouhl support a growth of wcMxUiestroying fungi at least as gurwl
as that on the malt extmct agar, and which at the same time would lie
cf>miM>se<l of as simple constituent'^ as |K»ssiblc. The medium could
then U» <iuplicat<Ml at any time, by any inv<»stigator. pn)viile<l the ch<»mi-
cals UM»d wen» of the same standani of purity.
For .satisfactory growth of w(HM|-<l<»st roving fungi a culture miMiium
mii*<t have, in addition to certain simple inorganic salts, the n€H*e9««ar>'
coin|Mitind.^ to furnish lM)th nitrogen aiul carlwin in a f<*rm n^dily available
to the fungii**. .Vmmonium stilts, nitrates and a.^paragin or its salts have
1
Thf pr«'M<rit p:ip(*r it* otH> of four pr('|):in*d l>y the junior authoc (l*ieprr» to
parti:il fultilhiH'tit of r«M|uirftiiriitM for th«' d«'Kn'<* of I)<»('tor of I'hilofMiphy in thr
liuvrrMty tif W i.Hr«»njtin.
' l.:tt«'r t«*f<t^ in thiH l:il)or:itory indicati* th:it In'^f in (»f littlr or no advanta|[r.
in m:in> c:im*m it luiiiirHhat rcturdH thf growth of W(KNl-<i«*i4t roving fungi.
1917] . PiEPER, Humphrey and Agree: Synthetic Media 216
frequently been used in synthetic media as a source of nitrogen, while
various carbohydrates have been used as a source of carbon.
The first step in this work was to select a nutrient solution of inorganic
salts with di-ammonium phosphate as the source of nitrogen. This solu-
tion was used as the basis for testing the nutrient value of various carbo-
hydrates and consisted of the following:
Di-potassium phosphate (KSHPO4) 4 grams
Di-ammonium phosphate ( (NH4)2HP04). • 2 grams
Magnesium sulphate (MgS04.7H30) 2 grams
Agar-agar (powdered) 15 grams
Distilled water 1000 cc.
Varying concentrations of the following carbohydrates, usually 20 or
40 grains in a liter of the nutrient solution, were used: Lactose, maltose,
cane-sugar, galactose, glucose and glucosamin. Growth of the fungus*^
was compared with that on malt extract agar as a standard.
The organism grew fairly well in every case. Cane-sugar and glucose,
however, gave the most favorable indications, the growth on the cane-
sugar being slightly more favorable. Since, however, the growth might
be changed by using other nitrogen sources in addition to di-ammonium
phosphate, it was decided to continue the use of both sugars in further
work.
The next step was to obtain a nitrogen source more available to the
fungus than di-ammonium phosphate, so the following substances were
tested by adding 2 and 4 grams, respectively, to a liter of the nutrient
solution containing forty grams of cane-sugar: Asparagin, sodium aspara-
ginate, ammonium asparaginate, caffein, guanidin carbonate, glycin,
leucin, creatinin and betain. With 0.2 and 0.4 per cent of caffein and guani-
din carbonate, respectively, no growth of the fungus occurred, but in all
the other cases a fairly good development was secured. The best growth
was, without doubt, obtained with asparagin and its sodium and ammoni-
um salts. The fact that asparagin alone might be a source of available
carbon for fungi was considered, but experiments showed that practically
negative results were obtained in the absence of a sugar.
Glucose was next substituted for cane-sugar while using asparagin and
di-ammonium phosphate as a source of nitrogen. The character and
rate of growth was the same as that obtained with cane-sugar. It has
been shown* that not all fungi contain an enzyme which will hydrolyze
cane-sugar, hence the substitution of glucose would presumably be an
advantage, and cane-sugar was therefore discarded.
* Fames annosus was used in all the preliminary tests.
* Boeseken, J., and Waterman, H. Akad. We tense h. Amsterdam, 20: 548. 1911.
Abstract in Bot. Gaz. 69: 413. 1915.
216 PinTOPATHOLOGY (VoL. 7
On the ha^is of th««o cxix^rimcnta a culture medium of the following
composition was selected for further test:
Ctlucose (CJiijOi), powdered 40.00
Di-potassium phoHphate (KjHPOi) 4.00 Krams
Ajtparagin (C«Ht()|Xj) 4.00 gnuns
Di -ammonium phosphate ( (NH4)iHP04) 2.00 grmim
Mafcnesium sulphate (MgS04.7HsO) 2.00 Krams
( *alcium carlMmate (('aCO|) 0. 25 {(rmm
Calcium chloride (CaClj) 0. 10 gram
Afcar-agar (powdered) 15.00 grmms
Distilled water 1000.00 cc.
It is evident that thb< medium Is synthetic except for the agar-agar^
which is used lis a sohdifyinfc agent. Its nutritive properties for four im-
portant wckmIh lest roving fungi wen* determined. As a comparison, simi-
lar t<*sts were made on another medium differing only in the 8ul)«titution
of Witte's |)eptone for jusparagin and di-ammonium phosphate. Although
this latter medium is not strictly synthetic, still a definite grade and purity
of the {)eptone can 1k» obtained from a reliable source. For many pur-
pos4's such a medium might Im* found of considerable advantagt% as has
Imhmi shown by t<*sts on Fotnes (trirwsn:< and Fomen pinicola rt»|X)rte<l later.
In pn»paring the synthetic me<lium, the mixtun* was heatcnl in a 1.^
liter fljisk in a watiT bath to prevent charring. It was then pounnl into
test-tulK»s lis quickly as iHJSsible, and well stirred during the operation
to avoid losing any precipitate that was formed. The tul)es were then
pluggcMJ with cotton and sterilizcil in live steam (1(K)®(\) without pressure
for thirty minutes on thrcH* succi'ssive <lays. A very slight pnn'ipitation
was f(nmd in the lM)ttom of the tulx^s after .*<terilizat ion. This was pn>l>-
ably a mixture of cal(*ium and magnesium phosphat4*s and car)N>nat4^.
\> small amounts of calcium salts s<H»m to incn»a.M» the vigor of the gmwth,
it is advi.sable to n^tain this s<Mliment by thoroughly shaking the tulM^si
U'ftjre (xmring into jx^tri disln^s.
The WitteV |><>ptone me<liuiii wjis pn»pan»d in the .**ame manner :is tlie
synthetic* and wa> <M»m|K)s<Ml of the following sul)stanc*f»s:
(iliiroHi* •( MiiiOci. piiwdcrrd 40 (M) fniinis
I)i-pf>taHHiuiii plio«4phati> (KilllH),) 4 00 grama
Witti-M |M>ptoiii* 4 00 grams
Mafcix'Miiiii Hulph:it<> MkS< >4.7li;< >) 2 00 grams
< 'ulciufii rarlHin:it(> (':i( '( >] i 0 25 gram
< ':ilriurii rhl«»ritl«' (*:i<'l: 0 10 gram
VK^tr-aicnr {Hiwdcrcd ■ 15 00 grams
Dintitlrd H.itor 1000 (K) cc.
' ( 'iill<»i<ial ;*ilirii* :i*'n\ ha** Imi'Ii ummI hy Horiif* wnrkrrs ill plai*e uf agar-agar and
might U> «if advaritaKf here l»ut the writrrM huvr not eX|M*rimented with it.
1917] PiEPER, Humphrey and Agree: Synthetic Media 217
The malt extract medium used for comparison was made as previously
indicated. It was tubed and sterilized in the same manner as the others.
All these media were tested with four wood-destroying fungi. The
tables give the growth of these at 25*^0. over varying periods up to twenty-
eight days.
From the results it is seen that the synthetic culture medium which
produced the best results gives a good growth and compares favorably
with the malt-extract medium. Perhaps by continued cultivation the
fungi may gradually become adapted to the new medium, giving then
better results than were obtained in this investigation.
The Witters peptone medium is especially good for the growth of Forties
pinicola and Fomes annosus. With Lenzites sepiaria, and especially with
Siereum frustulosuniy it gave less favorable results than the other two.
By additional work it may be possible to improve the synthetic medium
further so that it will give a better and more uniform growth than here
described. Such a medium would be very valuable both for toxicity
work and general physiological experimentation with wood-destroying
fungi.
In addition to the media for which the formulae are given in the pres-
ent paper sixteen other preparations were tried. The proportions given
are for 1000 cc. distilled water and 1.5 per cent agar. None of these were
as satisfactory as those reported above.
1. Cane sugar, 40 grams; ammonium asparaginate, 4 grams; magne-
sium sulfate, 2 grams.
Gave thick but slow growth for Fomes pinicola; wMi Fomes annosus
gave thin growth, covering the plate in two weeks.
2. Fraenkel and Voges' Solution.^
Gave verv poor growth.
3. Fermi's Culture-Fluid. «
Gave poor thin growth; plate not covered in three weeks.
4. Uschinsky's Solution.®
Gave poor thin growth; plate not covered in thre(i weeks.
5. Modified Uschinsky's Solution.*
Gave very poor growth.
6. Hasselbring's Solution.^
7. Cane sugar, 40 grams; glycerin, 40 grams; asparagin, 4 grams; mag-
nesium sulfate, 2 grams; di-potassium phosphate, 4 grams; di-ammonium
phosphate, 4 grams.
Gave best growth for Fomes annosus; thin growth for Fomes pinicola;
thick but slow growth for Fomes applanatuSy Lenzites sepiaria and Siereum
jrustulosum.
* See E. F. Smith, Bacteria in relation to plant diseases, Vol. I, p. 197, 1905.
"* Glucose, 1 gram; ammonium nitrate, 1 gram; di-potassium phosphate, 0.5 gram;
magnesium sulfate, 0.25 gram.
218
Phttopatholoot
(Vol. 7
TABLE 1
Radial growth in millimeUrs of Pome* annas tu in pariouM media at iS^C,
MEDIUM
NUMBBB or
rBTBt-DIUI
BADIAL OBOWTB OT rCMmVB W
MXLLMMWtmwm
7d«JB
lOd^ys
M dAf*
Synthetic <
Peptone . <
Malt-<»xtract |
1
r
2
2'
3
3'
21-23
23-24
23-24
24-26
•24-25
»V-28
30-34
31-35
35^37
33^39
29-34
30-35
40-12*
40-i8*
43-44*
40-42*
40-43*
41-45*
Character of growth:
Synthetic. Slightly more fliifTy and slightly leas dense than on mmlt-
extract, although the color was the same.
Peptone. Alx)ut the same as on synthetic medium but not quite as denat.
Malt-extract. Dense, creamy, fluffy growth.
The growth of Fomen annosus on synthetic medium and peptone medium was
tested for acidity with litmus and found to give a slight acid reaction after fourteen
days growth.
•Surface of medium entirely covered.
TABLK 3
Radial growlh in milUmrtrrs of Fomrn pinirola in variotix media at tS*C.
MBOIL'M
I
pEmi-Diaa
RADIAL OHOWm OT Pt'NUt'S IH MnXIMBTBBS
14 fUys ! ?1 dar*
limyn
10 <Uy»
Synthetic
1
1'
24-28
24-2t5
2 21-22
l*rpt«iiie
•>'
2<>-32
27-a3
33-35
30-35
Surface
covered
Surface
ctivrred
31-32 Surface
covere<l ,
Surface ■
cnvennl
31-32
Mnlt-4»\tr:irt
3
JfV 27 ! 32-:« I Surface
coveriMl
32-.'M : Surface
2.7 27
covered
Character of growth:
Synth*' tii'. Thill iirid Htriatni. rotor i«:tme as on malt -extract medium
iVptoiiv Wry fluffy and juMt .hm di*niH' ns on malt -extract medium
.Mttlt-fxtrart. White. denHe. fluffy growth.
1917] PiEPER, Humphrey and Agree: Synthetic Media
219
table 3
Radial ffrowth in millimetera of Lemites sepiaria in variotts media at $S^C.
MKDXUIC
NUM-
BBBOF
PBTBI-
DI8B
RADIAL GROWTH OF FUNGXTS IK mU.imTBBS
7 days
10 days
14 daya
21 days
28 daya
*
1
21-22
30-31
33-37
Surface
Synthetic
1'
22-23
'31-32
37-38
covered
Surface
k
covered
*
2
17-20
2&-30
34-35
37-39
Surface
Peptone -
2'
16-20
29-31
35-36
36-39
covered
Surface
b
covered
*
3
17-18
27-n30
33-35
Surface
Malt-extract -
3'
17-20
29-30
36-38
covered
Surface
k
covered
Character of growth:
' Synthetic. Margin of growth thin with much sub-growth; more fluffy
towards center than on malt-extract; color not as dark as
on malt-extract medium.
Peptone. Growth poor in appearance; color same as on malt-extract
medium.
Malt-extract. Fairly dense; dirty brown color.
table 4
Radial growth in millimeters of Stereum frustidosum in various media cU B5^C,
MEDIUM
NUM-
BER OF
PETRI-
DISH
RADIAL GROWTH Ol
' FUNGUS IN MILLIMETERS
7 days
10 days
14 days
21 days
28 days
»
1
9-12
14r-16
29-33
Surface
Sjmthetic •
V
10-11
15-19
28-34
covered
Surface
k
covered
Peptone <
2
4r-5
5-6
&-9
9-10
10-11
2'
7-«
9-11
10-13
12-13
13-14
»
3
11-16
21-22
26-28
30-36
Surface
Malt-extract i
3'
13-14
22-23
27-29
36-37
covered
Surface
k
•
covered
Character of growth:
Synthetic. Very fluffy, dense and creamy; slightly more brown in color
than on malt-extract medium.
Peptone. Poor growth; deep orange-brown color.
Malt-extract. Fluffy growth; slight yellow tint.
220 Phytopathology |Vol. 7
8. DifTrrs from No. 7 in substitution of 40 gmnis Kalactogo for th«*
vnm* suj[;ar.
(lavi* inurh slowcT growth than No. 7; platr not (\\uiv covitcmI in thnt-
WiM'ks.
S), Differs from No. 7 in substitution of 40 ^ranis lactose for th«- ran*-
sUKur.
( lavr slow growth, not as rocmI as No. 8; platr not cov(T<hI in thn*** \v«i*kv
10. Canr sug:ir, 40 ^^rains; glycin, 4 ^ranis.; (li-])otassiuni phosphate.
4 j^rams; nia^n<^iuni sulfati', 2 Rraujs.
(lave frmyi] jjjrowth; almost as rapid as on nialtn'xtract agar, but ap|x-ar-
anr<' not as j^ood as on No. 1.
11. DifTrrs from No. 10 in substitution of 4 grams rrratinin for th«-
glyrin.
(•a VI' fair growth: platr covircd in thriM* wrcks.
12. DitTiTs from No. 10 in thf sul)stitution of 4 grams, caflfrin for th«-
glyrin.
No growth in thn-t* wrrks.
Mi. l)iffrr> from No. 10 in th«' Mib«^tit\ition of 4 grams guani<lin rar-
l>onatr for thr glyciii.
(lavr no growth in three weeks.
14. Cant' sugar. 20 grams; glucosamin, 4 grams; (li-]><»tassiuni phi**-
phate. 4 grams; magn« vjum sulfate. 2 grams.
Fairly good growth, but less deii^r than nn maItHxtra<'t agar; plate
<M)vere<l in two wim k*^.
lo. (Ilueosr, 2.") grams; d and l-leuein, 4gram>; tli-|M»tassium phos]>hat«*,
4 grams; magnesiinn sulfate. 2 grams.
Platr eovrn-d in twn wrrks but growth thiiuur than on No. 10.
Hi. Laeto>e. 'A2 gran'>; brtain. 4 grams ;di-iM)tassium phosphatr. I Kni!ii«»;
inagin simn sulfate. 2 grams.
(Irowth fair; plate not cpiite eovered in thn^e weeks.
LirKKATlKK
.\s praeti<-;dl\ all of th<' literature dealing with synthetie eultun* ni(Nii:i.
a> far a> tlu* wrifi'r- have invest igati'd, ha^^ n^ferenee to its ada|)tabilit\
to tlu' growth of baet«Tia or mohls and as thex' organisms ap|):in*ntl\
H'aet to the media in a diffennt manner than the wtMKl-4h»>t roving llyni**-
nomyi-etes ufN>n which thi'^ stu<ly was eondu<*ted no l»ibliograp)i\ i^*
ap|M'ndtMl.
I\\ K>in;\iin\> IN rouKsi l*ArnnLn(;v. HiHKAr OK Plant Inoimhy
In ( uui'KHMioN Wirn thk ToiiKsr Piionrrrs Labokatoky.
Maoismn. \Vim onsin
PHYTOPATHOLOGICAL NOTES
Apple scab on the twigs. Does the apple scab {Venturia pomi (Fr.)
Wint.) pass the winter on the twigs, which thus become a source of in-
fection? This is a question frequently presented to the American plant
pathologists. The presence of the organism in the twigs has been re-
ported by a number of workers, but the first satisfactory^ data bearing
on the 'probabilities of this being a source of infection wei-e presented by
Morse and Darrow in 1913.^ They gav(? a very satisfactory review of
the literature of the subject, which will therefore be omitted in this
discussion.
In the latter part of April, 1916, the attention of the writers was called
to a nimiber of diseased apple twigs from Freehold, Monmouth County,
New Jersey. A careful examination showed that the organism was
Venturia poniiy and that the conidia which were present in great numbers
were viable.
The twigs showed an abundance of infection for several inches and the
bark was split, thus giving them a very ragged appearance. Directly,
beneath the ruptured bark were dense masses of stromatic growth, which
protruded through the openings and produced an abundance of conidio-
spores, which were short and irregular and rested on still shorter, in fact
almost equilateral, irregular basal cells. The spores were typical and
agreed in shape and size with those obtained from other sources, and
grew readily in culture. We are unable to say whether these spores were
formed in the fall of 1915 or the spring of 1916, but it is very evident
that the organism survived the winter in the twigs.
The writers did not have an opi)ortunity to examine the trees, but
Mr. W. B. Duryee, Jr., the County Farm Demonstrator who sent the
twigs to the Station, reported the disease in abundance. The twigs sub-
mitted to us carried an abundance of the organism, and the infection
extended from the tip back some 12 to 15 inches, but was most severe
near the tip.
Many of our New Jersey apple growers cultivate their orchards so
* Morse, W. J. Spraying experiments and studies on certain apple diseases in
1913. Maine Agr. Exp. Sta. Bui. 223. 1914.
Morse, W. J. Spraying experiments and apple diseases. Maine Agr. Exp. Sta.
Bui. 252. 1916.
Morse, W. J. and Darrow, W. H. Is apple scab on young shoots a source of spring
infection? Phytopath. 3: 266-269. 1913.
222 PnYTOPATiioLoaY [Vol, 7
thoroughly that fallen leaves an» vm' scarce* in the spring of the y«-ar.
However, thc^> same orchards will some times pro<luce an abundance
of «lis4»ase<l fruit. Although we fully realize* that a s<»vere inffTtion may
rt»sult from a small source, we have for some time lK»en inclimHl to Ix^lieve
that there must Ik» some source of infection other than the asco>|M>n'!»
foniUMl on the leaves of the prec<*iling year.
Mkl. T. Cook aSd C\ A. Schwarze
On Uiciug an ether freezing microtome in warm atui damp ire/ither. MoMt
|M*rsoiis who use an ether fn^'zing microtome may rememU*r that during
the warm and humid davs of midsummer and earlv fall then* is likelv to
Ik* coiisitlt*ral)lc ditiiculty in freezing material, wlu*n at otlitT tinle^ littlf
or no difficulty i> cx|M*rienc(>d. During the very damp and rainy day^
of f»arly sumnuT (P.)H») much difficulty of this sort was ex|N*rienc<Ml in
the I^il>orat(»ry of Fon'st Pathology, Providence, K. I. At times it was
absolutely imiM>ssible to frcM^ze the pn*paration. This IcmI to sonn* «*x-
IM*rimenting on the part of Mr. N. <). Howard. CollalKirator. ami the
writer, whit-h liniilly n'sulted in oven*<»ming tin* main difficulty in a vcr>*
simple and effi<*ient maimer. The a|>paratus <*:)nsisted m<*n*ly of a widt^
mouthe<l. eight- or ten-ounce l>ottle containing anhydr(»us calcium chloride.
which was in.^MTted )N'twe(*n the pn'ssure tatik and tlu* atomizer. Thbi
•
lM>ttlt* was connect<Ml so that tlu* air from tlu* tank pri-vsed thn»ugh a glasei
IuIn* in the stop|M*r to the hott >m of the botth*. and tliencM* up thn>ugh
the mass of cah'ium chloride to another gla.ss tiilH* whi(*li conncct4Hl with
the atomizer bv means of a rubiNT tuU*.
The calcium <*lil(»ri<le w:is broken into small piec(\s and |»acked intii
the Uittle. but not .so tightly :is to pn*vent the long glass inlet tuU* U-inie
worke<l down through tlu* mass as the stop|)er w:is ins<*rt4*d. The nil»-
Iht stopiMT was tied .s<»cun*ly in place so as to pn*vent its lM»ing blown
out by an i'Xc«*s.h of air pressure.
.-\fter a few (Livs of oc<*:i>i(»nal u.s4* the calcium chloride usuallv ^h4»u<*«i
sign> of (leliqueMvme through absoiption of moi.stun* from the air. Wlten
thi.s deli(|uesn'nce lN*came rather pn»nounc4*d, tli<* <'alciinn cliloride %ia>
planMJ in a small "fry-pan" and heated until it wits again entirely dry .tnti
hard, .\fter it had c<K>h*<l sufficiently to handle it was bn»k(*n up into
small pi(M*(*s and put back in the Inittle whih* still warm. Tlie same
cah'ium ehloride. which cost l»ut a few <*ents when |Mirchas(H|. |i:ls non
N-eii in Use for nion* than six months. When the fnn*zing microtome
\\a.s Us4*i| M'veral time> each day the <'alcium chloride hail t4> Ih' t|rit*ii
al'out every wi*ek or t**n davs.
.\lt hough the apparatus df*MTilNMl may Ih* consiflert*<l lis a soniewh:it
erude affair, it has worktMl verv effiri(*ntlv for mon* than six m«inths and
1917] Phytopathological Notes 223
has given no indication that its efficiency would not continue indefinitely.
Without doubt a more finished apparatus of still greater efficiency would
result from using a regulation chemical dehydrating apparatus.
Although this method of manipulation usually prevents the formation
of snow on the imder side of the freezing disk — ^which delays freezing —
it does not always prevent it. Recent experiments by Mr. Howard
show that this difficulty can be prevented entirely by placing several
small lumps of anhydrous calcium chloride in the ether bottle, or, better,
in the ether can itself as soon as it is opened, and letting it stand for half
a day or a day before using. This withdraws the small amount of water
in the ether, which apparently is partly responsible for the formation of
the snow.
In using ether for freezing sections we have always found it necessary
to filter the ether before it reaches the atomizer and also to avoid using
rubber in contact with it. The ether intake tube in oiu* microtome has
an inside diameter of less than 2 mm., and filtering is very easily accom-
plished by thrusting a small wad of cotton into the end of the tube. This
wad of cotton also is extremely useful in regulating the supply of ether
going to the atomizer, as, with a Uttle experience, the supply can be in-
creased or decreased almost at will by using respectively a loose wad of
cotton or a compact one. Of course the cotton used for filtering should
be renewed whenever it shows any tendency to become clogged.
This note is offered for pubUcation with the thought that other workers
who have had similar difficulties in freezing material with ether might
like to know that such difficulties can be overcome so easily.
J. Franklin Collins
Note on Xyhria polymorpha and X. digitata. The recent article by
Fromme and Thomas^ on a root-rot disease of the apple in Virginia, in
which the causal organism is provisionally referred to some species of
Xylaria, may be further substantiated by the following observations.
In 1906, near Scottsburg, Indiana, the writer collected mature speci-
mens of Xylaria polymorpha from diseased areas in living roots of a four-
years-old apple tree of the variety Winesap. The following year this
tree died and was pulled up. The conidial stage of the fimgus was after-
wards noted on the diseased roots. On October 5, 1908, in the same or-
chard Xylaria digitata was collected from the roots of a six-years-old pear
tree which had died from some unknown cause. In the writer's herbarium
are two other collections of Xylaria digitata, made at Priest River, Idaho,
from decayed areas in living roots of Populus trichocarpa and Crataegus
1 Science n. s. 46: 93. 1917.
224 Phytopathology (Vol, 7
douglam. The roots of the former wen* partially deca^'ed, but not in
the same part, by Fomea applanatuSf ami those of the latter by a species
of Follies* ix*culiar to this trcK*.
Jamek K. Weir
Puccinia triticina Erikss, Leaf -rust of winter wheat causes damage in
Kansas, It is j^encTally e<)nsi<lenMl that the leaf-nist of wheat due to
Puccinia triticina Krikss. is not s<»rious enoiigh to cause any appreciable
damaf^e to the erop, at h>ast publieations indicate that an attack of leaf-
rust in May or June does not produce* any marked effect on the yield.
()l>w*r\'ations by the \\Titer th<' past season showcnl that the Ieaf-ni5t
in some fields in Kansas was verv abimdant, an<l that its occurrence wa.*
not confim^l to the folia^ce but that the "necks" of the wheat wen» xngor-
ouslv attacke<l bv this rust. Careful o!)serv'ations and examinatioa«« of
«mch fields showed that no othir factors could have lM»en res|)oaHible for
the p<H)r cjuality of the ^rain and the rc<luced yield. The yield of one
variety in partic\ilar, a pure line winter wheat ^own in Kansas and
called P 7(M), was re<hic<Ml 8K jxr cait, acconlinR to yiehl data fumishe<l
by th<» Deimrtnii nt of AKnnnnny. The fields showing the effect nwirt
were thosi' whi<-h had bi'^'n planted lat<*. It is thought that this is par-
tially resjxmsible for the lar^c percent ag*' of leaf-rust.
The iMTcentaKe of infection on the "necks" of the wheat, as e;^iniat<*d
•by the newly adoi)ted scale for estimating nist iMTcentafC^^s of the Officf
of Cereal Investigations, I)e|mrtment of ARriculture, was 10 to 25 per
cent, while the folia^* of thf alM»ve-mentionc»<l variety Renendly showwl
KM) JXT cent of infection.
It is iM'lievi'd bv the writer that the leaf-nist of winter wheat in Kansas
can under favorable conditions ca\Ls<* con.sidenible damage and that t<Mi
little stress has In^en Riven in literature to the imi>ortance of this rust.
L. K. Mku*hers
Earhj (liacwery of white ptne blister rust in the Vnitni States, Then*
has nHM*ntly come to the atU'ntion of the writer the fact that a siM»cinien
<if white pin«* blister ru>t was c(»llceted nii white pine iPinus strobus Linn, i
by Mr. Samui'lN. Haxt«T of Philadelphia, in April, HM^o. at a nurs«*r>' n^-ar
Philadflphia. A s«-arch of the (>orresi)ondence in thr files of the rnited
Stat«-*« I>4*partmrnt of AKri<'ultiin* cnrrolnirates this statement.
The »»p<rinMn whi<'h wa> sent t4i the TnitiHl States |)<*|mrtm«*nt of
AKrirultun- w;i> r»'f«TnM| to th** Mycoloj^ist, Mrs. F. W. Patters<in. f«»r
I \\v\f. J:iiii«- \{ Nfiti'H nti \%M<HlHli>itrii>iiiK fiiiifei which ftrow on Uith romfrmui
and 'Iff nlii'Mi- xrw^ I. Phyi<i|):itli. 4: J72. VA\.
1917] Phytopatuological Notes 225
examination, and pronounced "a Peridermium which causes what is called
a pine-blister rust."
A search in February, 1917, in the pathological collections of the Bureau
of Plant Industry failed to reveal the specimen. The letter from Dr. L. O.
Howard, dated April 22, 1905, referring the specimen to Dr. A. F. Woods,
has the notation on it, * White Pine, Peridermium on'* in the handwriting
of the Mycologist.
Since there is no reason to believe that the determination was incor-
rect, and since but a single species of Peridermium has ever been reported
as causing a blister rust on white pine, this specimen was in all probablity
Peridermium strobi Klebahn, the pine stage of Cronartium rihicola Fisher.
This record, then, antedates the finding by Stewart of the Cronartiiun
stage on currants at Geneva, New York, in 1906, and the hitherto earliest
record in this country of the Peridermium stage on pine discovered in
New York on June 8, 1909, and reported by Spaulding in 1909.
Roy G. Pierce
Needle rust on Pinus resinosa. In June, 1916, the writer saw at Sharon
Vermont, a very striking case of needle rust in a plantation of 10,000
trees of Pinus resinosa about 4| feet in height. Up to about 3 feet the
yellow spore bodies, although small, were so abimdant on the 1915 needles
as to be readily discernible 15 to 20 feet away, single needles bearing as
many as 20 pustules. Hedgcock has identified the rust as belonging
to the two species, Coleosporium solidaginis (Schw.) Thimi. and C. deHca-
tulum (Long) Hedge. & Long. The alternate hosts were abundant in
the plantation as well as beyond it. This seems to be a case of healthy
trees being brought to that locality and infected by the fitngi which were
already present on the herbaceous hosts. Because of the abundance
of the rust the writer thought it might be a serious matter to such small
trees. A second visit made in September, however, showed that the
needles were a healthy green color except for small dead spots where the
rust pustules were produced. It is possible that the damage may later
become more evident and this point will be determined.
Perley Spaulding
Notes on the distrihution of the bacterial disease of western wheairgrass}
Until recently the writer had not observed the bacterial disease of western
* O'Gara, P. J. A paper read before the meeting of the American Phytopathologi-
cal Society, Columbus, Ohio, December 28, 1915.
Abstract published in Phytopathology 6: 98-99. 1916.
Science n. s. 42: 616-617. 1915.
Phytopath. 6:341-349. 1916.
336 Phitopathology [Vol. 7
wheat-grass, Agropyrmi smithii Rydlt., causoil by Aplatwbacter Agropt/ri
O'Oara, 4iut!<kic of two cMunties within the atato of Utah. The disease
has been recently found, however, occurring on wetrtem wheat-grass in
throe widely wparated districts of Montana, namely, Lewis and (,'lark,
Broadwater, and IXht Loilge counties.
Ab noted in previous papers, this disease is most characteristic in that
the yellow bacterial oose is fuuml to cover the glumes of the inflorescence
and appears also in droplets of considerable slie on the outride of the
^umes and un the sheaths. It also causes Minu> dwar6ng of the plants
aa well aa a Ix'nding of the stem alwve the last int^-mode. When the
inflorescence is infected gemiinable see<ls arc not produced.
The fact that this disease has l>een found in widely separated distrieta
would indicate that it may be found wherever western wheat-grass grows.
The writer would appreciate u note from anyiine finding this disea.'<e, as
he is interested in establishing the ext^'nt of is distribution.
P. J. O'Gara
Thr occurrence of Cotletotrichum aolania^um O'Gara on eggplant. A note
cunreming this species of C'olletotrichum was publinhetl as an almtrsct
in Phytopiithologj-.' Later ii description of the orgiminni appeare<l in
Myi-<iloi;i:( ■ At tht- time of pliblishiiiK my i-urliiT nutts there was M>me
dimbl a- t.i \h>- |i:ir:i-iti-ni of thi- m-w -pc<'ii-s. Ciiltun-s wm- exchangiil
MiCiini (' .r A .li.»-:iM' i.f iln- iiiKltricn 111 rill hiiiiii- nf Irii-li piitAUi rnuard by ■
Ii<-» ^|.ii... ..t l'Ml|.-i..lii,'|iillti l'livrr>|):itli 4: tl» III I'll).
'•>(;, r, ]• .r \.» .fi,.,!... .,rr..|l.ir..tri.l,i,..i:.[i.l ni..r>in. Myr..l.>Ki» T: :to -II.
1917] Phytopathological Notes 227
with Dr. J. J. Taubenhaus, then at the Delaware Agriculture College,
who concluded that the organism is a species of CoUetotrichum and that
it has parasitic tendencies (oral communication).
During the summer of 1916 the writer had occasion to observe a field
of eggplants in which fully ninety per cent of the plants showed wilt.
At first it was supposed that the wilt was due to a Fusarium, noting only
the general appearance of the field. A careful examination of the plants,
however, indicated that Fusariiun was not present but that the roots
and stems were badly infected with the above-named organism. Interior
portions of infected roots and stems were taken under sterile conditions
and placed in culture tubes, where the organism fruited characteristically.
No other organism appeared in the cultures where the proper precautions
were taken. Even in mixed cultures, Fusariimi did not appear.
When the organism was studied as a root and stem parasite of the
potato, it was not suspected of being a serious wilt fungus, but in the case
of the eggplant it has been found to product a wilt disease similar to that
•produced by Fusarium. During the early growth of the plants no trouble
was noted, but about the time some of the earlier fruits began to mature,
the infected plants wilted. This condition shortly prevailed throughout
the entire field, producing an almost total loss.
The field in which these eggplants were grown had supported a stand
of potatoes the previous year and it was in this field of potatoes that the
writer first found the above-named organism. From specimens collected
in this field the original description was made.
This note is published in order to give notice of the economic character
of this species of CoUetotrichum.
' P. J. O'Gara
Personals. Mr. K. E. Quantz, formerly assistant plant pathologist at
the Virginia Experiment Station, has become plant pathologist to the
Brazilian Government, with headquarters at Rio de Janeiro.
Mr. H. E. Thomas has resigned his position as instructor in plant
pathology at the Virginia Polytechnic Institute to accept an appointment
as assistant pathologist at the Federal Experiment Station at Mayaguez,
Porto Rico.
Mr. Fred R. Jones, formerly a graduate student at the University of
Wisconsin, has been appointed to the position of pathologist in charge of
forage crop disease investigations. Bureau of Plant Industry', beginning
April 14, 1917.
Mr. Gustav A. Meckstroth, a student at Pennsylvania State College,
has accepted an appointment as scientific assistant in plant pathologj',
Office of Cotton, Truck and Forage Crop Disease Investigations, Depart-
ment of Agriculture, beginning May 1, 1917.
LITERATURE ON PLANT DISEASES*
Compiled bt Eunick R. Obbrlt, Librarian, Burbau op Plant Industry, amb
Florkncb p. Smith, Absibtant
February to March, 1917
Arthur, Joseph Charles. Uredinales of Porto Rico bsBod on collections by H. H.
^lietiel and E. W. Olive. Mycolofda 9: 55-104. F. 1917.
Bloletti, Frederic Theodore and Bonnet, Leon. Little-leaf of the vine. Jour. Agr.
Research 8: 381-398, 2 fig., pi. 89 02. Mr. 6, 1917.
"A disease of the vine, vyinfc in seriousness with oidium and phylloxera."
Cause unknown.
Blsby, G. R. The short -cycled Uromyces of North America. (Abstraei.) Phjrto-
pathology 7: 74. F. 1917.
Blake, Msurice A., Cook, MelvUle Thurston, and Schwsrze, Carl Alois. Studies on
peach yellows and little |>carh. (Abstract.) Phytopatholoay T: 76-77. F.
1917.
Block, W. S., and Ruth, W. A. Control of apple scab by bleaching powder. (Ab>
stract.) Phytopathology 7: 76. F. 1917.
Brooks, Charles, and Cooley, Jscquelln Smith. Jonathan spot. (Abstract.) Phy-
topathology 7: 76. F. 1917.
Tom|)eraturc relations of apple rot fungi. (Abstract.) Phjrto-
pathology 7: 76. F. 1917.
Brown, Nellie Adaless. A bacterial stem and leaf disease of lettuce. (Abstract.)
Phytopathology 7: 63. F. 1917.
Boiler, Arthur Henry Reglnsld. Black rust of wheat. Bui. Misc. Inform. Kew,
1917: 4K. F. 1917.
I'>ononiir 1<his due to m'hoat rust in North America.
Bnrkholder, W. H. Bean diseases in New York 8tate in 1916. (Abstract.) Phyto*
pathology 7: 61. F. 1917.
Dry rcN>t rot, Futiarium sp.; blight, BacUrium Pfuueoli.
Butler, Ormond Rourke. How to control the snapdragon rust.
FhiriHts* Kx. 4S: 353. F. 17. 1917.
Dusting with sulphur.
Bysri, Luther Psrrls. A nomatrMle diHcase of the dasheen and its control by hoi
wat<*r tn'tttmcnt. (.Mwtrart.) Phytopathology 7: 66. F. 1917.
Tylenchtui triliri on whrat. Phytopathology 7: 56-57. F. 1917.
' fWfttnntnc with this numbrr. it i* intrndvtl that this liat thmll includ* all r«f «f««MB l» Um
luf« ol plant lii t'mar*. bbtb Amn-lrmn and f«»f«lgn All forriga mrinekf publiabad MBO* Jmumtt I. tflft
• hirh itnif t-.i our Bttrtiti'in. «ill hr rntrrcii. wi that th« imirs may hm ulttmAlvly
from tlut (lat*
Ail aut>tor« ar«> "rfirl Ut loi'iprratr in makinc thv lt*t C(im(»l«.# by ft«ndittc Ibair
ibC r«>rrrviftn« aiiit aiMitmn*. atwl r«|imall> by rallinc ait4*titi<>n to mrht4>n<iua
nf rvBular |ii-irnaU iip|irifit« <»r r<irTM*|M»fMlFtirfl* •liuulil tie aililrrMd lu Miaa K. R
BiirvAU i4 IMabt loduMo. l*. .*< I>*p« Agrx , WMhit^ton. I). C.
1917] Literature on Plant Diseases 229
Carsner, Eubanks. Do the bacteria of angular leaf spot of cucumber overwinter
on the seed? (Abstract.) Phytopathology 7: 61-62. F. 1917.
Chivers, Arthur Houston. The injurious effects of tarvia fumes on vegetation.
Phytopathology 7: 32-36. F. 1917.
Bibliographical footnotes.
CoUey, Reginald Hunter. Discovery of internal telia produced by a species of Cron-
artium. Jour. Agr. Research 8: 329-332, pi. 88. F. 26, 1917.
Literature cited, p. 332.
Cronartium ribicola.
Mycelium of the white pine blister rust. (Abstract.) Phytopathology
7: 77. F. 1917.
Cronartium ribicola.
Pycnial scars, an important diagnostic character for the white pine blister
rust. (Abstract.) Phytopathology 7: 77. F. 1917.
Cromwell, Richard O. Fusarium-blight, or wilt disease, of the soybean. Jour.
Agr. Research 8: 421-440, 1 fig., pi. 95. Mr. 12, 1917.
Literature cited, p. 438-439.
Ftaarium tracheiphilum Smith.
Cuba. Secretaria de Agricultura, Comerdo y.Trabajo. Comlsion de Sanidad
Vegetal. La enfermedad del cocotero, y medios para su exterminio. Cuba
Sec. Agr. Com. y Trab. Com. San. Veg. Circ. 2, 4 p. S. 1916.
La enfermedad del platano y metodos para combatirla. Cuba Sec. Agr.
Com. y Trab. Com. San. Veg. Circ. 3, 4 p. S. 1916.
Dalbey, Nora £. Com disease caused by Phyllachora graminis. Phytopathology
7: 56-56, 1 fig. F. 1917.
Damell-Smlth, G. P. A disease of spring flowering bulbs. Agr. Gaz. N. S. Wales
88: 141-142. F. 1917.
Davis, W. H., and Johnson, Aaron Guy. The aecial stage of the red clover rust.
(Abstract.) Phytopathology 7: 75. F. 1917.
Uromycea fallens (Desm.) Kern.
Doran, William L. Controlling snapdragon rust. Value of copper and sulphur.
Florists' Ex. 48: 501. Mr. 8, 1917.
"Bordeaux has no effect on the rust of snapdragons, and will not control this
disease."
Doubt, Sarah L. The response of plants to illuminating gas. Bot. Gas. 63: 209-
224. 6 fig. Mr. 1917.
Literature cited, p. 224.
Durrell, W. L. Notes on curly dwarf symptoms on Irish potatoes. (Abstract.)
Phytopathology 7: 71. F. 1917.
Edgerton, Claude Wilbur. A Gnomonia on eggplant. (Abstract.) Phytopathology
7: 78. F. 1917.
Perhaps identical with Phyllosticta kortorum.
Sdson, Howard Austin, and Schreiner, Oswald. A malnutrition disease of the
Irish potato and its control. (Abstract.) Phytopathology 7: 70-71. F. 1917.
XUiott, John Asbuiy. A new parasitic slime mold suitable for class work. (Ab-
stract.) Phytopathology 7: 74. F. 1917.
^aulwetter, R. C. Dissemination of Bacterium Malvacearum. (Abstract.) Phy-
topathology 7: 64. F. 1917.
Dissemination of the angular leafspot of cotton. Jour. Agr. Research
8: 457-475, fold, chart. Mr. 19, 1917.
Literature cited, p. 473-475,
Physiology of bacterium malvacearum. Smith. South Carolina Agr. Elxpt.
Sta. ^th Ann. Rept. [1916]/16: 49-64. [1916.]
230 Phytopathology (Vol. 7
Pronime, Fred Denton, and Thomas, H. S. Spraying and dusting tomatoes. Vtr*
ginia Agr. Expt. Sta. Bui. 213, 14 p., 3 fig. 1916.
Fromme, Fred Denton. A Xylaria root-rot of the apple. (Abstract.) Phyto-
pathology 7: 77. F. 1917.
Gardener, Max William. Dissemination of the organism of cucumber anthracnoae.
(Abstract.) Phytopathology 7: 62-63. F. 1917.
Gilbert, William Williams. Virulence of different strains of Cladosporium eum-
merinum. (Abstract.) Phytopathology 7: 62. F. 1917.
Gladwin, F. S. and Reddick, Donald. Sulfuring Concord grapes to prevent pow*
dory mildew. (Abstract.) Phytopathology 7: 66. F. 1917.
Great Britain, Board of Agriculture. 8clen>tinia diseases. Jour. Bd. Agr. [Lon*
don) 2S: 109:>-1008. pi. 1917.
ScUrotinia sclerotiorum.
Varieties of imtatooK roMiHtant to wart disease. Jour. B<i. Agr. [London]
88: 1017. Ja. 1917.
These varieties are more fully deMrril>ed in Garden 81: 8-9. Ja. 6. 1917.
Hedgcock, George Grant, and Hunt, N. Rex. An alternate form for Coleosporium
helianthi. (Abstract. > Phytopathf»logy 7: 67-68. F. 1917.
Noten on some .H|H»cies of (.\>Ieosi>orium. (Abstract.) Phyto-
pathology 7: VtS. V. 1917.
The Peridermium l>elonging to Coleosporium ipomopv. (Abstract.'
Phytopathology 7: 67. F. 1917.
\ IVri<lermium U'longing to Colecwporium terebinthinace*. (.\b-
Htraet.) Phyt<»pathology 7: 67. F. 1917.
- Some new hosts for Colcos|N>rium inconspicuum. (Abstract.)
Phytopathology 7: 6H m. F. 1917.
Some new hosts for Coleosporium solidaginis. (Abstract.) Phyto*
path<»l<»gy 7: tlS. F. 1917.
Hedgcock, George Grant, and Long, William Henry. The aecial stage of C<»K
s|H)rium elephantop<uiis. (.Xhstract.) Phytopathology 7: 66-67. F. 1917.
Periderm turn came u m .
Hlggins, Bascombe Britt. A dis4*aM> of |N»ean catkins. Phytopathology 7:
2 fig. F. 1917.
Hongerford, Charles W. Pureinia graminis an wheat kernels and its relation to
sulHM'quent inff*ction. (Abstract.) Phytopathology 7: 73. F. 1917.
Jackson, Herbert Spencer. A six*eies of (*hrysomyxa new to North America. ;.\1>-
HtriMt.i Phyt<»pathology 7: 7«. F. 1917.
t >n Picra engelmannii.
Jagger, Ivan C. Tw<i transmissible mosuie diseaM*a of cucumbers. (Abstract ;
PhytiipHtholufcy 7: 61. V. 1917.
Jeffrey, J. W. MyHterimis vine «lis4*aM'. .Mo. Hul. Stati* Com. Hort. (Califomim)
6: 4in 117. N. P.Mr,
"It ilitfN Hot :ip|K*ar that w thontiigh examination of this malady has ercr
Ihm'Ii iii.-idf, hut exp«'rts have determined that the cause is not Phylloxera. Aha-
hi-ifii di-i-:i>f. fungus nr biM'teria."
Jehle, Robert Andrew. < 'h:ira<'terii*tirs of eitrus ranker and of the causal orgma*
f«iii i^Mi:irt Mul StHt<- IM:int Ii«l. Floriila 1: 'J4 'SI. illus. Ja. 1917.
P*t 'tdiifuitu'i* ritri.
Citru" I nrikiT irive!(tiKati4»MS at thr Klorifla tnipiral lalmratory. (Abstract >
Pli>t«.p.itho|Mj:v 7: .> :»9. F. 1917.
Johnson, Aaron Guy, ami Coerper, Florence M. .\ baeterial blight of soy beaa.
(Abstrart ) Phytopathiihtgy 7: t>5. F. 1917.
1917] Literature on Plant Diseasi^ 231
Johnson, Aaron Guy, and Hungerford, Charles W. Scolecotrichum graminis on
timothy, orchard grass, and other grasses. (Abstract.) Phytopathology 7:
69. F. 1917.
Johnson, M. O. The spraying of yellow pineapple plants on manganese soils with
iron snlphate solutions. Hawaii Agr. Expt. Sta. Press Bui. 51, 11 p., 4 fig.
D. 13. 1916.
Johnston, John Robert. Enfermedad de la rafz de la cafia de azticar. Pt. II. Ha-
cienda 12: 146-147. F. 1917.
History and cause of the rind disease of sugar cane. Jour. Bd. Comm. Agr.
Porto Rico 1: 17-46. pi. 1. 1917.
Bibliography, p. 43-45.
Melanconiufn sacchari,
Jones, Fred Reuel. The Pseudopeziza leaf spot diseases of alfalfa and red clover.
(Abstract.) Phytopathology 7: 70. F. 1917.
Jones, Lewis Ralph, and Bailey, Ernest. Frost necrosis of potato tubers. (Ab-
stract.) Phytopathology 7: 71-72. F. 1917.
Johnson, Aaron Guy, and Reddy, C. S. Bacteria of barley blight seed-borne.
(Abstract.) Phytopathology 7: 69. F. 1917.
Keitt, George Wannamaker. Peach scab and its control. U. S. Dept. Agr. Bui.
395, 66 p., 6 fig., 6 pi. (1 col.). 1917.
Literature cited, p. 64r-66.
Cladosporium carpophilum Thtim.
Second progress report on investigations of leaf spot of cherries and plums
in Wisconsin. (Abstract.) Phytopathology 7: 75-76. F. 1917.
Elrout, W. S. Bacterial diseases of celery. (Abstract.) Phytopathology 7: 64.
F. 1917.
Crown rot, Bacterium and Fusarium; crown rot wilt; bacterial heart wilt.
Wintering of Septoria petroselina var. Apii. (Abstract.) Phytopathology
7: 65. F. 1917.
Lathrop, Elbert C. The generation of aldehydes by Fusarium cubense. Phy-
topathology 7: 14r-16. F. 1917.
Lee, H. Atherton. A new bacterial citrus disease. Jour. Agr. Research 9: 1-8,
pi. A, 1-2. Ap. 2, 1917.
Bacterium citrarefcicienSy sp. nov.
Long, Wilham Henry. Five undescribed species of Ravenelja. Bot. Gaz. 61: 417- •
424. My. 1916.
Ravenelia roemeriancBf sp. nov.; R. morongiae, sp. nov., R. thornberiana,
sp. nov., R. reticulata, sp. nov., R. anntdataf sp. nov.
Investigations of the rotting of slash in Arkansas. U. S. Dept. Agr. Bui.
496, 14 p. 1917.
Oak slash: — Stereum rameale, S, umbrinum, S, versiforme, S. fasciatum
Shortleaf-pine slash: — Polystictus abietinus, Lenzites sepiaria.
None of the main fungi concerned in rotting either the oak or the pine slash
in Arkansas produce heart rots in living trees.
A preliminary report on the occurrence of western red-rot in Pinus ponde-
rosa. U. S. Dept. Agr. Bui. 490, 8 p. 1917.
McClintock, James A. Economic hosts of Sclerotinia libertiana in tidewater Vir-
ginia. (Abstract.) Phytopathology 7: 60. F. 1917.
Lima bean mosaic. (Abstract.) Phytopathology 7: 60-61. F. 1917.
Peanut-wilt caused by Sclerotium rolfsii. Jour. Agr. Research 8: 441-448,
pi. 96-97. Mr. 19, 1917.
232 Phytopathology (Vol. 7
McClliitock, Jamei A. Will SponKospora subtcrranea prove serious in VirgiBiA*
(Abstract.) Phytopathology 7: 72. F. 1917.
McCnbbin, W. A. Doen Cronartium ribicola wintor on the currant' PhjrtoiMitbol-
o|0' 7: 17-31. 1 fig. F. 1917.
Bibliography, p. 31.
\Vhit<^ pine blister rust on currants. Canad. Hort. 40: 34, 1 fig. F. 1917.
Mackle. D. B. Observations on the distribution of citrus canker. Philippiat
Agr. Uev. 9: 278-281, pi. 4. 1916.
Martin, William H. A sclerotium disease of peppers. (Abstract.) PhyiopatholoflQr
7: M. F. 1917.
Sclerotium bat alt cola.
Msi4, Pierre Marie. Chlorose toxique du mais, la s6cr6tion interne ei la rW stance
natun^llc df*8 v^g^'taux sup^ricurs aux intoxications et aux maladies paraai-
tain>M. (Nmipt. Hcnd. Soc. Biol. [Paris] 79: 1059-10($6. D. 1916.
Melchers, Leo Edward, and Dale, E. E. Black spot of popper. (Abstract.) Phy-
topathology 7: 03. F. 1917.
Alternaria sp.
Melhus, Irving E. Notes on nxmiac symptoms of Irish potatoes. Phyiopniholofcy
7: 71. F. 1917.
and Diehl, William. The development of the aecial stage of Nigredo on red
clover. (.\bRtra4t.i I*hytopathology 7: 70. F. 1917.
Merrill, J. H. Further data cm the relation between aphids and fire blight (Bacil-
lus amylovorus (Bur.) Trcv.). Jour. t>on. Knt. 10: 45-46. pi. 2, F. 1917.
Murphy, Paul A. The economic importance of mosaic of |>otato. Phytopathology
7: 72 73. F. 1917.
Seed |M>tato certification in Nova Scotia. (Abstract.) Phytopnthology
7: 72. F. 1917.
llorton, John Bitting Smith. Host limitations of Spctoria Lycopersici. (Abstract. )
Phytopathology 7: 65. F. 1917. '
Ilowell, William. Summary of notes on fungi and plant diseases. Agr. News.
{Barbados) 16: 14. Ja. 13. 1917.
Olive, Edgar William, and Whetzel, Herbert Hice. Kndophyllum-like rusU of
Porto Uico. Amer. Jour. Bot. 4: 44 .'>2. 3 pi. Ja. 1917.
Botryttrhiga Hipftocratctr \Vheti4*l Ar Olive, sp. nov.; EndophyUum eircum-
itrrifdum (Schw.) \yhetn*l A Olive, comb, nov.; Endophyllum W^ddiw (Earle)
Whetiel A Olive, comb, nov.; Endophyllum decoloratum (Schw.) Whetsel A
Olive, comb, nov.; Endophyllum Stocky tar phettr (Henn.) Whetsel 4 OliTe.
c<in»b. nov.; EndophylloitleM portoricenifiit Whelwl A Olive, sp. nov.
Osner, George A. Pn^Iiminary not4*s on a m*w leaf spot of cucumbers. (Abstract. )
Phytopathology 7: 62. F. 1917.
Patten, Charles G. < >rigin and development of hardy, blight-resisting pears. Minn.
Hort 46: 97 Uf2. Mr. 1917.
Pennington, Leigh Hunt. Boleti Hnd niycorhisa u|M>n f«»n*8t trees and an unusual
iii>ri>rliiia upon white oak. (.\l>tttr:irt. ) Phytopathology 7: 74. F. 1917.
Pierce, Roy Gifford. Albany c(»nfen*n(e on white |>ine blister rust. Phytopathol-
ogy 7: :a .v.. F. 1917.
Potter, Alden A., and Melchers, Leo Edwsrd. Kcological observations on I'siilago
Z. :». Ph>inp;»tholngy 7: TH 74. F. 1917.
Rand, Frederick Vernon. U-afiipot-rot of pmid lilien raumMl by Ilelicosponum
n>tiiph:uruni. Jour Agr. It«M-arch 8: 219 2:i2. pi. 67 70. F. 5. 1917.
1917] Literature on Plant Diseases 233
Ruikiii, WiUiam Howard. The control of white pine blister rust in small areas.
(Abstract.) Phytopathology 7: 58. F. 1917.
The penetration of foreign substances introduced into trees. Phytopathol-
ogy 7: 5^13, 1 fig. F. 1917.
Ravn, Frederik K^^lpln. Jens Ludwig Jensen (1836-1904). Phytopathology 7:
1-4, port. F. 1917.
Reed, George Matthew. Phytopathological survey of the trees and shrubs of Pros-
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work. Brooklyn Bot. Gard. Rec. 6: 14-20. Ja. 1917.
Rhoads, Arthur S. Some new or little known hosts for wood-destroying fungi.
Phytopathology 7: 46-48. F. 1917.
Rosenbaum, Joseph. Studies of the genus Phytophthora. Jour. Agr. Research
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Literature cited, p. 273-276.
and ShapofvaloY, Michael. Strains of Rhizoctonia. (Abstract.) Phyto-
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Nematospora Lycopersici n. sp.
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Storage rot fungi.
234 Phytopathology (Vol. 7
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P.M7
PHYTOPATHOLOGY
VOLUME VII NUMBER 4
AUGUST, 1917
THE CUPRAMMONIUM WASHES
Their Preparation, Biological Properties, and Application
O. Butler
With Plates III to X
A number of cuprammoniums have been recommended as fungicides,
but, despite the fact that they form, or it would be more accurate to say,
could be made to form, clear solutions and leave, on drying, inconspicu-
ous spots on the foliage, no single one has sufficiently met the desiderata
of practice to become generally employed. Practice demands that a
fungicide be non-toxic to the sprayed plant at the strength at' which it is
most efficient and effective, but the cuprammoniums have the reputa-
tion of being unduly toxic and of not being as effective though more
efficient than Bordeaux mixtures.
The toxicity of the cuprammoniums has been ascribed to soluble cop-
per (Millardet^ being notably a proponent of this view), to excess am-
monia, and to ammonium sulphate. The prevailing opinion regarding
the ineffectiveness of the cuprammoniums is based on their behavior
relatively to Bordeaux mixtures which, in fact, is comparing solutions
weak in copper with mixtures considerably stronger. Hawkins,* for
instance, compared a cuprammonium containing 0.053 per cent copper
with Bordeaux mixture containing 0.18 to 0.25 per cent copper. The
question may, therefore, be asked, are any or is any one of the cuprammon-
iums worthy of being retained amongst the fungicides? In order to satis-
factorily answer this question it will be necessary to study: (1) The com-
position and preparation of the different washes. (2) The relative
toxicity of the different washes, and the conditions affecting the same.
* Millardet, A., and Gayon, U. Les divers proc6d68 de traitement du mildiou
par les composes cuivreux. Journ. Art. prat. 1: 729. 1887.
• Hawkins, L. A. Grape -spraying experiments in Michigan in 1909. U. S.
Dept. Agr. Circ. 65. 1909.
236 -Phytopathology [Vol. 7
(3) The relative efficiency and effectiveness of the cuprammoDiumft.
This we will now proceed to do.
I. CHEMISTRY AND METHODS OF PREPARATION OF THE CUPRAMMON'U-MS
In the preparation of the cuprammonium washes met with in practice
either animoniiim hydroxid or ammonium carbonate are employerl an
the solvent, the solute being, in the case of the former solvent, either
copper tuminf^s, i.e., metallic copper, copper sulphate^ the basic <*<»pper
carlwnatc malachite (CuCOj. Cu(()H)x), or the basic copper carlK>DAt«
of Burgunily mixture (2 CuCX),. 3 ('u(()H),)» and in the case of the latter
the copper salts mentioned, together with basic copper sulphate and cu-
pranunonium sulphate (CuS()4. 4 NHj. HjO). With the exception of
the copper sulphatisammonia wash which Ls a cuprammonium sulphate,
all the other waslu»s preparwl with ammonium hydroxid produce cupnuu-
moniums of very similar, if not identic composition and fonii a well
charact<»riz(Hl group; similarly the wjisIh^ prepared with aimncmium
carl)onato an* also compositionally so ne^irly alike as to form a well lie-
fincnl gn)up. The* fonner ar(» cuprammonium hydrati^s, the latter cupram-
monium carlN)nat<^. Since (^uprammoiiium sulphate was the first in-
tnxluced of the cuj)ramm<)nium washes and the cuprammonium hydrates
were introduccHl prior to the cupranunonium carlMmates, it will lie possible
lK>th to n*tain a group distribution and consider the M^veral fungicid<<9( in
chn>nological ord("ir.
,4. Cuprammonium Kulphate wafihen
CopiHT sulphnlv and ammonia. The copfx^r sulphate and ainmunia
wash (ejiu c^lc^stcO was the first iiitnKluc(Kl of the cuprammonium fungi-
cidfw and still n^nuiins. taking the world at large, the l)ost known. It 15
tlu* most easily pn<^|)anxl and at the same time the most unstable, even
when sufficient amnumium hydroxid Ls (iinployini to give a clear h>Iii-
tion. The fungicide was intHNluccnl by AudoynaucH in 18S5, but the
original fonnula which is as follows:
Cupriciiulphntr 1
Aiiitiumiuiii hydroxiti ^p. gr. 9() 0 769* by volume
WiiXt'T to 100
* li4Mifi)r(l. Dukf of. uimI Pick«*nnf(. S. l*. Wotmrn Kxperimental Fruit Fi
H«pt U: S4i. IIMO
' Aij«loyiiniiii. A. Ia* miUlioti c*t Im roinp<iA<'t< cupriqucs. Progrteagr. el nt.
IKK'.
* Kfiuivnii-nt of animoniiim hydrntr up. gr. 92, 1 part by volume called for in cht
origpiml foriiiula.
1917]
Butler: The Cuprammonium Washes
237
proved injurious to vegetation and has suffered ftiore or less marked
modifications at the hands of subsequent writers as will be seen from a
perusal of table 1.
TABLE 1
Strength oj copper sulphate and ammonia wash and ratio cupric sulphate: ammonium
hydroxid recommended by different authors
CUfRIC SULPHATK
STRBKOTH IN CUPRIC SULPHATB
AMMONIUM HTDROZID
SP. OR. 0.90
per cent
■
0.26
1:1.50
0.50
1:0.76
0.50
1:1.06
0.50
1:1.70
0.50
1:1.14
1.00
1:0.73
1.00
1:1.06
1.00
1:1.46
The formulae given in table 1 do not necessarily represent the composi-
tion of the fungicide when put in service, however, for we find authors
recommending that it be allowed to stand after being made for a few
hours, half a day or even several days* so that the excess of ammonium
hydroxid may pass off. Nor do the formulae permit, except in one in-
stance, i.e., in the case of the wash containing 1 per cent cupric sulphate
and a ratio cupric sulphate-ammonium hydroxid of 1: 1.46, of the fung -
cide being applied as a perfectly clear solution which is essential if tlie
copper is to be deposited on the sprayed foliage in the proper physical
and chemical states. Millardet^ long ago pointed out tliat Audoynaud's
formula was in this respect defective and increased the amount of am-
monium hydroxid so as to give a ratio of 1:1.53 instead of 1:0.769.
The ratio cupric sulphate ammonimn hydroxid required to give a clear
1 per cent solution (and Millardet's figures are correct) has to be greatly
increased in order to prevent a precipitate forming when washes contain-
ing less than 1 per cent cupric sulphate are employed. The Duke of
Bedford and Pickering* have shown that the amount of ammonium hy-
droxid required increases very considerably with the dilution and the
data presented in table 2 confirms this view. But while the data given
in the table show a very considerable increase in the amount of ammonium
* Viala, P. Les maladies de la vigne, ed. 3, p. 143. 1893.
^ Millardet, A., and David, E. Essais comparatifs de divers proc^ds de traite-
ment du mildiou. Compte-rendu Congr^ nationale viticole, Bordeaux. Appendix,
60. 1886.
• Woburn Exp. Fruit Farm Rept. 11: 18. 1910.
238
Phytopathology
(Vol. 7
TABLE 2
Amount of ammonium hydrorid required }o give clear nolutionM of copper MtUpkmie
and ammonia of different ntrengthn in cupric sulphate together unth the corrt»pomd-'
ing ration cupric sulphate: ammonium hydroxide and percentages of ammonia
VrRKNUTH IN
per rtnl
I 0
0 '>
0 2
0 1
AMOI'Nr AMMONlim
■TDSOXID MBQt'lKBD TO
flIVB rUBAR ftOLUTIONi
per cent
15
1 0
0 8
0 7
II4TIO
CUPKIC •rLTMATB
AMMONII M HTDMOXIO
1:15
1:20
1:4 0
1:7 0
IK NHa tn»m-Anri
OMLT
0 386
0 2.SS
0 251
0 179
'The* data given under the heading amount of ammonium or ammonium rarliunjit^.
as thecaacmay Iw, required to give a clear 8olution indicatea the amount of the Milt
required to give a clear solution for a ixTicMl of time of not leiM than two houm in
a closed veatM*!.
hydroxid n»lativo to tho copper, it also brings out equally clearly that
the al)solute amount of anunonia* in solution decreases as the conr(*nt ra-
tion of the copfMT is lowennl. It may also )>e well to add that onitiK
to the extn^mc; volatility of ammcmium hydroxid the fi^reit given in the
table will have to Ih» inrnMistMl at temp<Tatures much alnive 20**(\ :uk1
conversi»ly may 1k» somewhat <h»rnjas(Ml for tomiKTatures l)elow 2iY*i\
The rop|NT sulphate and ammonia wash may l)e most conveniently
prepare<l by adding the ammonium hydn>xid nH)uiro<l to a strong si>lu*
tion of copfMT sulphate and diluting imnuHliately after the pnH*ipitat«
first fonn<»tl has dissolvtui, though the wash may Ih» satisfactorily pn»-
pannl even when the n^lative dilution of the salts varies f^ithin wxilc
limits. It is g(*4ierally considi^n^l that the copper and anunonia wash
(and the other cupranunoniums) are In^st pn^panni with soft water Mncp
the solutions tin*. d(H*ompos4Ml by hard water. The amount of copper
pnH'i(utate<i wliftn hani water b( usimI is, however, entirely m^Ugible in
practi(*<» lis nuiy Ih» judgcMl frofu table 3.** *
TABI.K 3
Am4>unt of cttpiHT itrrripitated from thi mpfur »ulphnte and amm4ntia womK hy ir«i.'rrt
iff difftrt ut dtgntM of htirdttrMx
IM>I tt'K or WATRR
H%iibMiciM or I
MrTHH- OKllRKU ,
corrva
raBrirrr»Ti»
(*ily «if liord«>.*iu\
(!nvoii ykvW
-.Yi 0
92 0
0 OOONt
0 ocMm
*The ii'Tiu afiiitionia in \imh\ to denote Nil, unlefut the context indicatm tlie
connotation Ml .Oil
'* Mdlardrt. A . and tiayon. V. Id** divert* prtWuirii de traitement du mtlcfaou
par Ira comfHMM'i* ruivreux. Journ. .\gr. prat. 1: 7.Ti. 1KK7.
1917] Butler: The Cuprammonium Washes 239
When ammonium hydroxid is slowly added to a strong solution of
cupric sulphate a precipitate of basic cupric sulphate (CUSO4. 2 Cu(OH)i)
is thrown down which dissolves in an excess of the reagent forming a
deep blue solution, the copper being then in the form CUSO4. 3 NHj. 2
HjO" or, according to the more commonly accepted view CUSO4. 4 NHs.
H2O a salt which decomposes readily on volatilization of ammonia or on
dilution, the copper' being precipitated as a basic sulphate. When the
fungicide dries upon foUage the copper is deposited as a basic sulphate
mixed with a Uttle ammonium sulphate in the ratio of 1:0.35; in other
words only a very small amount of the latter salt can be present even
when a 1 per cent solution is sprayed on foliage as will be clearly seen
from a study of the reactions involved which are as follows:"
(1) 3 CUSO4 5 H2O + 4 NH4 OH = CUSO4. 2 Cu(0H)2 + 2 (NH4)2S04
+ 15 HjO.
(2) CuSO.. 2 Cu(OH), + 2 (NHOjSO, + ;„ "' "T = 3 (CuSO*. 4
in excess
NHs.HjO) + 9 HtO.
(3) 3 (CUSO4. 4 NH3.H,0) + H2O = CUSO4. 2 Cu(0H)2 + 2 (NH4)
SO4 + 8 NH,
B, Cuprammonium hydrate washes
A cuprammonimn hydrate is formed when metallic copper, cupric
oxide, cupric hydrate, malachite, or the basic cupric carbonate of Bur-
gundy mixture are dissolved in ammonium hydroxid, and washes have
been used in practice prepared from copper and all the copper salts men-
tioned with the exceptions of cupric oxid and hydrate. All the cupram-
monium hydroxid washes decompose on dilution or on volatilization of
ammonia with formation of cupric hydrate, the copper also being de-
posited in this form when the fungicides dry spontaneously on foliage.
The cuprammonium hydrate washes are more stable than cuprammonium
sulphate.
Copper and ammonia wash. The copper and ammonia wash, or
Schweizer's reagent, was first introduced as a fungicide by Bellot des
Miniferes in 1887" but despite the fact that the results he obtained are
said to have been highly satisfactory it is practically unknown in the
literature.
For the preparation of the copper and ammonia wash a very large
amount of ammonium hydroxid is required and the copper must be acted
" Bedford, Duke of, and Pickering, S. U. Woburn Experimental Fruit Farm
Kept. 11: 17. 1910.
** Chester, F. D. The copper fungicides. Journ. Myc. 6: 23. 1891.
" Bellot des Mini^res, H. Ammoniure de cuivre et parasites de la vigne. 1887.
240 Phytopathology (Vol. 7
on in thc^ firesenco of air or traces of aniinoniuni salti). Bellot de» Mm-
i^res employed the fonner method and prepared a stock solution whirh
was diluted at time of use so as to contain 0.25 to 0.75 per cent metallic
copper, i.e., the copper equivalent of a 1 to 3 per cent Bordeaux mix-
ture. The stock solution was made as follows:
Copper turnings 1
Ammonia, up. gr. 0.9 .' 119 by volume
A stock solution prepare<l in the manner indicated can l)e dilutc<l with
water to 0.0^^17 per cent copper without a precipitate forming within a
pericxl of two hours. At this dilution, however, the solution c<mt2iins
0.97 pcT cent ammonia which, as a glance at table 4 will show, is much
higher than in the other cuprammoniums of (H]uivalent stn^ngth in i-oi^
per. The fact coupled with the difficulties incident to the prep^iration
of the st<K»k solution hjis sc»riously militate<i against the emplojnnent of the
fungicide »n practice, the advantage's incident to its iLHt\ i.e., an ailht*!*ivi*-
ness wpial to the copper sulphate and ammonia wash'* and U»sser injuri-
ousness to the grape,** l)eing not sufficiently comp<»nsator>' to outwf*i|^h
the dmwbacks.
TABI.K 4
Amount of ammonia riquiud to ffiir t Irar solutions of the ruprammttnium ir«i«A/«
irhrn containing 0J)!M7 ftrr cent mrtallic copjter
NAMC or ruNoiriDr.
( *op|MT-nmnioniii
(\)|i|M>r ((ulphiit<'«nnim(iniA
(*np|M*r xiilphato- iinirnoniuni rHrl>onat4*
Malnrhite-amnionia
Hiirgimdy niixture-ammc»nia
Malafhitr-ntiimonium rarbonatr .
Hiiricuiiily niixtun*-aninioniuni carhonati*
4MOr»T NHi BB4Jt »«•
TO OIYB % iLBkB
ptr cent
e 9772
0 179
0 OI9
0 GMM
• ,ViO
0 047
0 aiH
Hunjundy mixture and ammonia. This the ftarlii»st copjier carkNinate
and ammonia wash employe<l in practi(*e wt&s intnNluce<l by Patrigi^in
in 1KS7** and coiL*<i.»*ts simply in dissolving th<' ba»*ic coppcT carlNmate
of Burgundy mixtun* din'i'tly in the mothcT li(iuor by moans of am-
monium hydn)xi<i. The w:ish Ls us<h| to some extent in practice and
'• Fo*\. (1. ("ourr* roriiplct dr Vitirultiin* ed. 4. 577.
" 1 1»^\. <1. TXw iMiiiu*. p. .'>7H.
^* I'ntrigroii. (t. Nouvraux pHN'^l^ di> tr»it4*m(*nt dii mildiou. Journ. m^-
prat. 1; s-sj. ish7.
1917]
Butler: The Cuprammonixjm Washes
241
is known in the United States under the name of modified eau celeste.
The formula originally proposed was as follows:
parte
Copper sulphate 1
Sodium carbonate 1.5
Ammonia sp. gr. 0.9 0.769 by volume
Water to *. 100 "
Patrigeon's formula has not suffered any marked modifications at the
hands of the various authors who mention it as will be seen from table 5.
TABLE 5
Strength of Burgundy mixture-ammonia and corresponding ratios copper sulphate
sodium carbonatCf and copper suLphaie ammonium hydroxid recommended by dif-
ferent authors
BTRENQTH IN CX7PRIC SULPHATE
COPPER SULPHATE
RATIO
SODIUM CARBONATE
COPPER SULPHATE
RATIO
AMMONIUM HTDROXID
per cent
0.25
1:1.20
:0.83
0.30
1:1.25
:0.83
0.50
1:1.25
:0.83
0.50
1:1.25
1.04
1.0
1:1
:0.73
1.1
1:1.25
:0.78
The amount of ammonia called for in the various formulae for the
preparation of modified eau celeste is never sufficient to give clear solu-
tions. The basic carbonate of copper of Burgundy mixture, separated
from the mother liquor by decantation is, however, readily soluble in
ammonium hydroxid, as may be gathered from table 6.
TABLE 6
Amount of ammonium hydroxid required to give clear solutions of modified eau cileste
STRENGTH IN COPPER SULPHATE
AMOUNT OP AMMONIUM HTDROXID
REQUIRED TO GIVE CLEAR
SOLUTION
STRENGTH OF SOLUTION IN NH|
per cent
per cent
per cent
1.00
4
1.029
0.50
3
0.772
0.25
2.5
0.642
0.10
1.4
0.360
Burgundy mixture dissolves in ammonimn hydroxid, forming cupram-
monimn hydrate and is not a mixture as Chester^^ believed of cupram-
" Chester, F. D. Report of the Mycologist. Delaware Agr. Exp. Sta. Rept.
4:68. 1891.
242 Phytopathology (Vol. 7
monium hydrate and carbonate. Modified eau o^leste does not fcive
jw. to malachite on decomposing thus indicating absence of cupram-
monium carlx>nate. When the wash dries upon sprayed foliage the cop-
per is deposited as an hydrate, mixe<l with some sodium sulphate and
bicarl>onate unless the copper carlx)nate is separated from the mother
lic]uor and levigated l>efore l)eing di«<solved as is sometimes recommended.
Copper carbonate and ammonia. The copper carbonate and ammonia
wash, or ammoniacal copper carl)onate was introduced by Gasline'' and
in American fungicide literature is the cupranunonium most commonly
met. with.
Malachite dLssolvos in ammonium hydroxid forming a solution which,
according to the Duke of Be<lfonl and Pickering,'* consists mainly of
cupranunonium cariM)nate, while (-hester*® Lh of the opinion that Inrth
cupranunonium carbonate and hydrate are forme<l. As, however, the
malachite ammonia wiish dcM^s not deposit malachite on standing. I
incline* to the view that the copper is pres4»nt solely as cuprammonium
hv<inite.
Malnchit<» (iissolv(\s sparingly in anunoniurn hydroxi<l, the total amount
entering into solution lK»ing l(»ss in a strong than a weak concentrati<in
of ammonia. P<»iuiy** for instance, found that a 42.()8 per cent ammonium
hvdn).xid dLssolviHl. imt gram weight of ammonia, 0.01329 grams of nHv
tallic co|)ixT while !ind<»r similar conditions a 21.38 percent solution din-
soIvchI 0.3132 grams and a 3.20 |K>r cent solution 1.063 grams metallic
i'opiMT n*sjH»<»tiv(»ly. The.s<» results, unfortunately for practice, can not
Im> obtaine<l by dis.Holviiig a given (plant ity of malacliite in the suitable
corr<*s|)on<ling strength of ammonium hydn)xid. In order to obtain the
maxinnun s<»lvent action it Ls nec<»ssar>' to us<» a ver>' large excesw of mala-
chite, **even five fold or more" which int roduc(*s ob vioas difRculties t hat
can not Im» tununl cxct»pt in a v(Ty empirical an<l unsatwfactorj' manner.
The malachite-anunonia wjish was, {is I have aln*ady indicat<*<l, pn>-
|M>MMi by (fiistine whos4» formula w:is jis follows:
Mnlarhitr . . 0 M
.Viiiiiiniiia Kp i^r. n.lN) ... 0 76U
Watrr to .100
Th(* alK>v<* fonnula hit** Imkmi iiiorr or U'ss m<Nlifi(Hl at the hands of
subMH|U(*nt writers, as will W\ .»<4»<»n from table 7.
'* (iaMtiiK'. i\. Ijiiploi <lij f*:irlM»ii:it4* aiitiiioiiianil di* cuivre contfc* Ip pm>ni>*
P|Hira I*r«»([ agr i-t vit. 8: 1SS7.
»\\i»lMirii K\p Fruit I ami \U\>x 11: JI !«»I(I
• I III* ««arii«'. p ♦">>*.
■> rititix (' I. 'I'ljf p-«paratiiin nf :tninii»iiiaral miIiiIioii of ropprr carlmiuilr.
Dc'laman- Aier l.xp ."^la Mill 22: .'> iyi:{.
1917]
Butler: The Cuprammonium Washes
243
TABLE 7
Strength of malachite^ammonia wash and corresponding ratioh malachite: ammonium
hydroxid recommended by different authors
STRKNGTH IN MALACHITE
MALACHITE
BATIO
AMMONIUM HTDROXID
per cent
0.045
22.2
0.046
5.5
0.060
12.8
0.075
11.3
0.078
:10
0.078
:13.2
0.086
:10
0.093
:8.7
0.093
:5.5
0.097
:10
0.097
:13.2
0.100
:1.23
0.100
:11
0.125
:8
0.200
:8
Neither in the original formula nor any of the subsequent ones that
have come to my knowledge is the ratio malachite-ammonium hydroxid
such as to insure complete dissolution of the copper salt. In order to
dissolve the malachite completely, at least within a reasonable time, the
ratio must be increased to 1 : 30. It is therefore clear that, as usually
prepared, the wash either contains less copper than the formulae call
for, or if the undissolved malachite is incorporated in the wash then the
copper will be placed on the plants partly in the form of malachite and
partly as a copper hydrate. In order to obviate these difficulties I have
employed a stock solution prepared as follows:
parte
Malachite t ? 1
Ammonia sp. gr. 90 30 by volume
Water 20 by volume
In preparing the stock solution water must be used as otherwise the
solution would prove unstable, decomposing %vith formation of cupric
oxid. A stock solution containing as little as five parts water may be
prepared and probably more water than the formula calls for could be
used, though it will be perfectly obvious that there is no object in making
a stock solution unduly dilute. In fact, ceteris paribus highly concen-
trated solutions are to be preferred.
The stock solution as above prepared is quite stable and may be diluted
very considerably without decomposing, but contains more ammonia than
244
Phytopathology
(Vol. 7
TABLE 8
Amount of ammonium hydroxid required to ffive cleat nolulions oj maiarhiU*
ammonium hydroxid
•TIIKKOTH IN TEBMi OF
corpftM arLniATB
prr rent
1
0 5
0 25
0 10
AMOUNT AMMONIUM HTOKOXIS
MBQUIRBO TO UtVK CLBAB
•OLUTION
ce.
26 5
13 '25
6 {\2
2 65
•TBBNOTV or •OLTTION IN NHf
6 82
3 41
1 70
0.6S2
mcNiified eau celeste for (Hjual p<»rcoiitttg«s of copper (Cu) as may l>e Hsen
by comparing tahlo 8 with table (>. Then^ is, therefore, no just ifieat ion.
since sodium sulpliate and biearlM)nate are not injurioii:^ at the conrcm*
trations at which the wjtsh may l)e employed in practice, in the preference
aocordi^d the malachite-anunonia wash, as the (*opper (M*curs in the same
fonn in Inith. The difTen»nce in favor of modifi(Ml iiau c^Uwte is n<it only
markcMi. however, in the fonniihu> I have us<»d, but Is also favorable to
the latter wh(»n w<» comparer \\u\ formulm^ of authors. Taking the ejc-
tn»nM^ met with we find tlie n«ults shown in table 9 wliich an* even
mon^ favonible to modified eau c^U^ste than in the case of my fonnulae.
And when we consider the (^ost of the unit copper employiHl (exclasive
of lalK)r) in nuHlifi(Hl eau c^l(*ste and malachite-ammfuiia we obtain a
ratio of 1:3 wliich is so siKnifi(*ant as to n^iuin^ no comment.
TABLK 9
Extrrmr jterrrutagrtt of ammnuin nut with in thr Utrmidae of author* for the ffrefiaro-'
tiou of numiifitd rait n'lrHtt * A ) and mat ac hit i -ammonia \H\ renpertirtty
•TRrMiTM Of HOLI^ION IN
TKBMM or rorrcH
■I Lrn «TC
per ttnt
0 J.")
1 1
nTKKM
iTH
OF
MiLi-ri«».M
iM
NH*
A
\
B
prr rrnt
ptremni
0 051
:
0 554
0 (»73
0 141
0 joi;
0 707
CI JIM
2 S2S
t'. t'liprntntnitnium cnr^Httuitf wnshrs
< 'upniiniiKiiiiuni carlNiiiatc \\:i^h(•^ an* foriiuNl when cupric sulphate,
ni:il:i<*)iiti*. rupraniiiifinMim >ulph:it<* '('uSn4. -| Nils. H3O) and the Imisic
cupn«' *:irlM»iiai<* nf HurKuiidv tnixtun' an* ilirv-olvnl in runmoniuni car-
ImukiIi MMf< t); MI4. NH,<'n, . Till- \\a>h«-^ an» ver>' sUble. the
nMi>t «t:tii|i- III till* riipraiiiiit«>iiiuni fun^icidi*^. only decom|M)sinK slowly
1917]
Butler: The Cuprammonium Washes
245
on long standing with formation of malachite. When the washes diy
on sprayed foliage the copper is deposited as a carbonate.
Cuprammonium suiphcUe-amm^mium carbonate. This wash was in-
troduced by the United States Department of Agriculture in 1890**
under the name of mixture number 5 but has never been used to any ex-
tent in practice as it was not found to possess a lesser toxicity than other
cuprammoniums and is not economical to prepare. The formula origi-
nally proposed was as follows:
porta
Ammoniated copper sulphate (Cuprammonium sulphate) 0.21
Ammoniiun carbonate 0. 12
Water to 100
Malachite-ammonium carbonate. This fungicide was introduced by
Chester** in 1891 as a substitute for the malachite-ammonia wash, but
despite manifest advantages has been but Uttle used in practice. Ches-
ter's formula is as follows:
parts
Malachite 0.052-0.058
Anmionium carbonate 0.27 -0.31
Water to 100
The original formula has been modified to some extent by subsequent
writers as i^ shown in table 10.
table 10
Strength oj malachite-ammonium carbonate wash and corresponding ratios malachite:
ammonium carbonate recommended by different authors
MALACHITK
STRSNOTH IN 1IA.LACHITE
AMMONIXTll CARBONATK
percent
0.039
1:6
0.046
1:5.3
0.052
1:5.19
0.058
1:5.34
0.066
1:3.3
0.093
1:5.3
Malachite dissolves fairly readily in ammonium carbonate, carbon
dioxid and a little ammonia being evolved during the reaction, but the
reaction is not sufficiently rapid nor the conditions under which it takes
place such as to permit the preparation of the fungicide as required. A
stock solution is necessary and may be conveniently prepared as follows:
« United States Department of Agriculture Rept. 1890: 402.
» Delaware Agr. Exp. Sta. Rept. 4: 71. 1891.
246
Phytopathology
[Vol. 7
Malachite 1
Ammonium carbonate (hard)** 3
Water 30
Place the malachite in a suitable non-metal vessel, add the amnionium
carbonate in small pieces, and then the water. Warm gently and mb
soon as effervescence bepns remove from flame and stir. Let Ktand a
few minutes, place back on flame and continue as before until on wann*
ing fi^ntly no further effervescence takes place. The vessel should thiro
be closed and set aside until the malachite has completely disHolved,
should it not already have done so. The stock solution prepared as aljove
will withstand marked dilution without a further addition of anunonium
earl)onate I>eing require<l at least within the ranfi^e of concentration in
coppcT that I liave used, as will lx» seen from table 11.
TABLE 11
Amount of ammonium carbonate {hard) required to give clear ttolutionn of malarhitf
ammonium carbonate
n'RKNOTM IM
AMOt'NT AMMOMCM
(-\HBONATE NKgriHKb
TO ur\'K A (LJCAM
IM»LrTION
HALAcnrrK
STRCffOTII or IMILC-
MAi.4rHnrB
AMMONIt'M C'AKBONAnC
TioM m NH*
prr rtmt
prr ctnt
^tretmi
0 .V)
1 5
1:3
0 476
0 JO
0 f)
1:3
0 19U
0 10
0 3
1:3
0 oas
0 o:>
0 15
1:3
0 047
Copper Hulphaie and ammonium carbonate. The coppt;r sulphate and
anunonium car)K)nate wash known in Anu^rican literature as Johnson'^
mixtun; has lHM»n but little uschI in pnictic(» an<l Is but ran»ly mentione^i
by writers on the funfci<'idi*s. The wiu<h was first dtwcrilKid in 18!U by
J(>hns<»ir* who profKiMHl the following formula for its preparation:
(NjpJMT Htllphfld'
Ariiiiioniuin riirlK»iiiiti> (hard)
(ir Ariifiioriiiiin rnrlNtrmtr 'Htift )
WfttiT to . . .
0
1
0
21)
0
23
1(10
'* Aiiiiiiofiiiiiii rarUtiiatc diM^oiii^MiW!* (»ri cxpoHurc tf> nir ami in prrparinfc th<* M«K*k
iu>ltitii>n th«' atiiiiunt of thr Halt ii^mmI will have (o U> incroamHl unirm it ia in prr-
f«M tly hard tranNlurrnt platrH. \\ hrn romplft<>ly df'<*oniiM»m>d ammonium rarlKinjite
ocriirH lis an o|Ktc|uc ihiwiIit and wh<*n in thin romlition the amount railed for in
tli«- fi*niitl]a ^h(lllld U* dollhlrd.
" Jnliiit^oii, S \V Noll' l»y ihf Dirertor. ConniM'tirut A^r. Kxp. Sla. Kept.
1890:113 iy«l.
1917]
Butler: The Cuprammonixtm Washes
247
Johnson's mixture is, next to the copper sulphate and ammonia wash
(eau celeste), the most easily prepared of all the cuprammoniums. When
anmioniimi carbonate is added to a strong solution of cupric sulphate a
precipitate is first formed accompanied by effervescence due to the lib-
eration of carbon dioxid which on a further addition of ammoniimi car-
bonate promptly and completely dissolves even in the cold.
Johnson's mixture forms very stable solutions and, as a glance at table
12 will show, is for equivalent of copper compositionally identical with
malachite-ammonium carbonate.
TABLE 12
Amount of ammanitan carbonate hard and soft required to give clear solutions of John'
son's mixture and corresponding ratios cupric sulphate: ammonium carbonate
■TBXNOTH IN
AMOUMT AMMONIUM CABBONATE
BSQUIBND TO OIVB A CLBAR
■oxjUTIom
CUPRIC SULPHATE
STRENGTH
or SOLUTION
IN NHt^
PHATB
AMMONIUM CARBONATE^
Hard
Soft
P€reerU
percent
percent
percent
1.00
1.56
2.40
1:1.56
0.495
0.50
0.78
1.20
1:1.56
0.247
0.20
0.31
0.48
1:1.56
0.099
0.10
0.15
•
0.24
1:1.56
0.049
^ Refers to solutions prepared with undecomposed ammonium carbonate.
Burgundy mixture-amnumium carbonate. Burgundy mixture may bo
readily dissolved in ammonium carbonate 3delding a cuprammonium
very similar to those obtained with malachite, copper sulphate, or cupram-
moniiun sulphate as will be seen by a glance at table 13.
TABLE 13
Amount of ammonium carbonate required to give clear solution of Burgundy mixture-
ammonium carbofKUe and corresponding ratios Burgundy mixture: ammonium car-
boncUe
BTRENOTH IN COPPER
SULPHATE
AMOUNT AMMONIUM
CARBONATE (HARD)
REQUIRED TO OnTE A
CLEAB SOLUTION
BUBGUNDT MDCTUBB
BATIO
AMMONIUM CARBONATE
STBENGTH OP SOLU-
TION IN NHs
percent
percent
percent
1.00
1.20
1:1.20
0.380
0.50
0.60
1:1.20
0.190
0.25
0.30
1:1.20
0.095
0.10
0.12
1:1.20
0.038
The data given in table 13 are for a Burgundy mixture in which the
ratio copper sulphate sodium carbonate (crys.) was 1: 1.84, and the pre-
cipitate was separated from the mother liquor before dissolving in the
248 Phytopatholoot [Vol. 7
ammonium carbonate. In practice, however, the precipitate would not
need to be separated from the mother liquor, since the salts therein con-
tained are not injurious^ at the concentrations at which the wash may
be employed in practice.
As a result of our study of the properties and preparation of the cupram-
moniiun washes we may conclude:
1. The cuprammonium carbonate washes are the most stable and for
strengths in metallic copper of 0.14 per cent or less require less ammonia
to give clear solutionis than the washes prepared with ammonium hydroxid.
2. The copper salts dissolved in ammonium carbonate 3rield washes
in which the active principle, i.e., the copper is in the form of a carbonate.
3. Metallic copper, malachite, and the basic carbonate of Burgundy
mixture form cuprainnionium hydrates when dissolved in ammonium
hydroxid modifie<i eau c^K^e requiring pvir equivalent of copper the
least amount of solvent.
4. Cupric sulphate forms with ammonium hydroxid a cupraiiunonium
sulphate which Is the least stable of the cuprammonium fungici<ics.
though less anunoiiia b< required to give a clear solution than in the case
of the niprammoniiun hydrates.
II. RELATIVK TOXICITY OF THE CUPRAlfMO.VIUM WASHES
The cuprammonium washes may l)e toxic to the sprayed plant: {D
Between the time of application and time of drying; (2) after dr>nng
owing to dissolution of the contained copper on weathering; or (3) the
injur>- pnxluced, if any, may l)e the result of the additive effect of 1 and
2. Thon^ lin'i no other possibilities.
A. Effect of the cuprammoniuffm on planta between the time of applicaiian
and the time of drying
It will Ix) at once evident that if the cuprammoniums nuule with ammoo-
iuiii hy<in>xid owe their toxicity to the presence of anunonia, the toxic
action, owing to the rapid dis.sipalion of ammonia in the interim lietwec^
time of application when its con(*ent ration will Ina highest and time of
d<*>i(vation when its conci^ntnition H-ill \)o zero, must and can only take
phi«*<* during the ilrying of the .^pray. Ammonium carbonate bt lev
vol.'itilc than junmoniuni hydroxid, but nevertheless decomposes readily
on rx|H»sure to air and has vani.shc<l when the wiishes of which it is a com-
ponent have dritni u|K)n the foliage, hen<*<* injur>' due to the presence of
ainin«iiii:i in ainn ionium ciu'lMmate can also only In^ produced in the
1917]
Butler: The Cuprammonium Washes
249
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to
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250 Phytopathology [Vol. 7
*
interim between time of application and time of drying. If the toxic
action is due, on the other hand, to noluble copper or ammonium sulphate
whenever this Halt occurs, then the injury resulting may be produced dur-
ing the time of drying and also at such sul)sequent times as the fungicide
is wetted by meteoric water. The rate at which the cuprammoaiums
dry upon sprayed foliage will, it may therefore be presumed, have a very
marked effect on the degree of injury produced. And in fact experimental
evidence fully confirms the presumption as is shown in table 14 and plates
III and IV.
The data presented in the above table show how significant was Audoy-
naud's reconmiendation that the copper sulphate and ammonia wash
be applic<l during dr>% wann weather and fully justifies Bourcart*.^'^
statement that scorching '*is especially to he feared when eau c^Ie:<te i^
applied during moist weather and, coni^c<}uently, when it dries slowly on
the leaves;** but that on the other hand ''when the spray is applied <lur-
ing the hot days of summer, this fault disappears entirely, and eau c*elf^te
possesses nothing but advantages.*' The data presented in the table
show conclusively that slow drying is much more injurious than quirk
drying, the difference l)eing ver>' striking indeed in the ca**e of malachite-
ammonia and malachite-ammonium carbonate containing 0.28 per cent
and 0.14 per cent copper.
While the data prosente<l in table 14 show that whenever a cuprain-
monium is toxic it is invariably more injurious when drieil slowly im^
spective of the plant spmyetl, it does not give us any ver>' definite infor-
mation regarding the cause of the deleteriousness of these washes. The
data do not supiN)rt conclusively either the view that aimnonia is the
toxic agent, or the view that the toxicity is due to soluble copper since
were but one of tlu»sc^ conijKments tin* sole cause of the injur>' protluccti a
certain proportionality would exist differing only in degree in different
plants lM!tw(H'n a given strength of the toxic Kul»stance and the resulting
mjur>', but a considenition of table 14 shows that no such relation exists
in cither c]is4*. On the other han<l, the data clearly show that ammonium
sulphate do«rs not |K)ssi*ss luiy niark<Mlly injurious properties. Neither
an* the malachite wa*<hes shown to In' K^s injurious than those maiie
fn»in cupric sulphate and auuiionia. a fact worthy of S4*rious consiiiera-
tion in view of the* general abandonment of the latUT cm account of sup-
fjosiil gr«iter toxicity.
Hut siiKM* nu individual coin]Minent of the cupranimoniuni wa*«h«*s is
apiMin-ntly inr x# the mn^r of the toxie action pnKluce<l in the inteival
lx*twe( n thf- applieatjnn of th«> fungirid«> and its desiiraticm it will U*
• liourrnrt. K I^-h iiiii1:m!i»'< «!«f plant h. .'JTii. Pnrirt. W^U).
1917) Bdtlbb: The Cuprahhoniuh Washes 251
necessary for us to study their behavior when applied separately in order
to interpret the data ^ven in table 14, and to the action of soluble copper
we may well devote attention first.
The only suitable copper salt to use in studyii^ the effect of soluble
copper when applied in the form of & spray is cupric sulphate and since
the acid radicals of the copper salts are not in themselves injurious*" the
toxicity of cupric sulphate may be considered as due to the copper. Sol-
uble copper is, as is well known, extremely toxic to vegetation and we
would, therefore, expect that the injury produced by an application of
cupric sulphate would be the greater the slower the spray dried on the
foliage, and the data presented in table 15, fully confirm this expectation,
and the illustration shown in plate V is no less emphatic.
TABLE IS
Effect of quiek and tlow drying on the loxieity of cupric tulphate eijtreued in per cent
of injury
> The beading, used also ia plate viii, means that the percentage strength of the
solution ie given in terms of milligrams iastead of fractions of a gram.
The data presented in table 15 and shown graphically in plate VIII
indicate that the tomato, Oxalis, bean and cauliflower are much more
sensitive to soluble copper than either the Coleus or the Pelargonium.
The tomato, bean, Oxalis and cauliflower respond in a somewhat simitar
mamier to soluble copper, the curves for quick and slow drjii^ being
" Bedord, Duke of, and Pickering, S. U. Woburn Experimental Fruit Fann
Kept. 11: 1910.
Clark, J. F. Oa the toxic properties of some copper compouods with special
refereoce to bordeaux mixture. Bot. Gaz. 33: 39. 1902.
Hawkins, L. A. The influence of calcium, magneBium and potaaalum nitrates
upon the toxicity of certain heavy metals towards fungus spores. Physiological
Researches 1: S7. 1913-16.
252
Phytopathology
[Vol. 7
ver>' nearly parallel at the higher concentrations. In the case of the cauli-
flower which is very sensitive to soluble copper the curves are nearly
proximate, while in the case of the tomato, which is much more tolerant,
they are distant. The curves for the Oxalis and bean are very similar
in character, and resemble those of the tomato, though the bean is more
sensitive to soluble copper than the Oxalis and the Oxalis less resistant
than the tc^mato. In the most highly resistant plants studied, the Pelaricrv
nium and C'oleus, the curves show marked similarity, the to:;icity on slow
dr>'ing increa«<ing a little more rapidly in the former than in the latter.
It will also be noticed that while in the more resistant plants ((*o!eua«
Pelargonium) the toxicity is practically proportional to the concentratioo
in the less resistant plants proportionality ceases to exist as scMin as the
threshold of toxicity is appn ached, the decrease l>eing much more rapid
than the data for the higher cxincentrations would lead one to anticipate.
The practical importance of these facts will escape no one.
TABI.F 16
Effect of hIow nrui quick druiug on the toxicity of ammtmi um hti<iroTid ■/». gr. 90,
eTjrreiitted in fnr cent of injury
\ ANIKTT mtV
HOW
DNIKD
nTHt NUTH (»r AOLrTION IN N Hi
lU 21) I 5 14 i 4 II 6.^7 2 (A I.2H
l^'T iwr ! |«rr I prr . iht jtrr
wilt ct'Ui ' <i*i»! I rent , o'lit r««iit
1 I
102
frnt
0 51
Tomnto
CiiiiliflowtT
'(^uirkly
.Slowly
Quirk ly
Slowlv
(^uirkly
Slowly
(^Uiirkly
Slow I V
per
inr ^ per \ prr per , per
cent • cent rent , cent . ctnt ! rwnt
IS
7ti
I
VA) !
7.'»
29
50
X\ i I 21 i
2:< I Iti
17 :i:>
10
I 25
I '^^
; 4ii
0 <
5
per
CtHl
I
s
12
34
O
a
93$
rrflkl
fw%S
0
5
22
0
U
. 10
Thr t<»xi<*ity of :iiiiiiH»niuiii hy(ln>xid h:is Ih»<mi ronsi<len»d higli by all
writers who have h<»ld the t<»nct that the toxi<*ity of the copp<»r ami am-
monia wastu's was due to th«* amiiionium hydroxid though MillanlH
|)oure«|, he tells us. stnmg aiiimonium hydroxid on the foliagf^ of the grafie
without pHMlucing injur>'. a result >tn>ngly (Mintradicting the prevailing
vi(*w. and the data pn>ent<sl in table Ki are not on analysis favorabh* to it.
A glanrc at the table revrals that like the cuprannnoniums and ruphc
sulphate* aniinoniuin hydroxid even though highly volatile is nmch UMkn*
t4»\ir when dri«»<l slowly and the illustnition given in plate VI sh*»ws
verv "^trikinglv what marked differences mav Ik* obtaineit. The tfata
al*4» sh<»w that, of all the plants us4<l, the tomato is the most sensitive
1917]
Butler: The Cuprammonium Washes
253
to ammonium hydroxid while the Coleus, bean and cauliflower follow in
increasing order of resistance. Now if ammonium hydroxid is primarily
the cause of the toxicity of the cuprammoniums of which it is a compo-
nent part the relative resistance of the plants when sprayed with the
copper sulphate and ammonia wash or the malachite-ammonia wash
should remain imchanged. In reality we find that when the former is
employed the sensitiveness of the plants is in decreasing order as follows:
Tomato, bean, Coleus;** and when the latter is used that it is as follows:
Tomato, Coleus, bean. The evidence is therefore clear that anunoniiun
hydroxid is not the primary cause of the toxicity of the cuprammoniums.
The toxicity of ammonium hydroxid is of course due to the ammonia
(NH3) it contains and it would therefore be expected that ammonium
carbonate which has been used in the preparation of cuprammonium
washes would also prove injurious since while less volatile and alkaline
than the former it nevertheless decomposes rapidly on exposure to the
air with liberation of ammonia and carbon dioxid. And in fact experi-
mentation shows ammonium carbonate to be much more toxic than
usually supposed as will appear from a consideration of table 17.
TABLE 17
Effect of slow and quick drying on the toxicity of ammonium carbonate, expressed in
per cent of injury
▼ ABIETT U8XD
Coleus "j
Tomato <
Cauliflower
Bean
STRENGTH OP SOLUTION IN NHj
HOW
DRISD
1.90
0.05
0.47
0.23
percent
percent
percent
percent
per cent
per cent
per cent
per cent
Quickly
40
2
0
0
Slowly
100
81
0
0
Quickly
40
15
5
0
Slowly
100
70
50
20
Quickly
75
29
10
7
Slowly
92
42
13
6
Quickly
1
0
0
Slowly
8
0
0
The data presented in table 17 and more strikingly illustrated in the
graphs (plates IX and X) show that within the limits of toxicity ammonium
carbonate is much more injurious per equivalent of ammonia than am-
monium hydroxid and it is rather diflicult to understand how it could
ever have come to be considered less noxious than the former. As would
be expected from the nature of the salt, the toxicity of ammonium car-
*<» The cauliflower was not used in the experiments on the toxicity of the cupram-
moniums.
264 Phytopathology [Vol.7
bonate when dried slowly is much more marked than when it is dried
quickly. The illustration (plate VII, fig. 1), will show what striking
effects may be obtained.
The data show that of all the plants studied the tomato is the most
sensitive to ammoniimi carbonate, the susceptibility of the other plants
being in order of increasing resistance as follows: Coleus, cauliflower,
bean. Similarly in the case of ammonium hydroxid, the relative
susceptibility of the plants is in increasing order as follows: Tomato,
Coleus, bean, cauliflower. The relative toxicity of ammonium hydroxid
and ammonium carbonate is clearly shown in plates IX and X. An in-
spection of the graphs on these plates shows that the toxicity of am-
monium carbonate, whether dried quickly or slowly, increases much more
rapidly with the concentration than does the toxicity of ammonium
hydroxid, the curves for the former indicating proportionality between
concentration and toxicity while those for the latter resemble those given
by copper sulphate. A consideration of plate IX shows that ammonium
carbonate is more toxic to the bean and cauliflower irrespective of the
rate at which dried than ammonium hydroxid. From plate X, on the
other hand, we gather that in the case of the tomato ammonium carbon-
ate is less toxic between 0 and 0.95 per cent ammonia than ammonium
hydroxid when dried quickly, but is invariably more toxic when dried
slowly; in the case of the Coleus ammonium carbonate dried quickly is
less toxic between 0 and 1.02 per cent ammonia than ammonium hy-
droxid and, when dried slowly, less toxic between 0 and 0.51 per cent.
Ammonium sulphate has been considered by certain writers as the
toxic agent of the cuprammoniiuns in which it occurs and the copper
sulphate and ammonia wash has suffered relegation in consequence of
this view. Pearson,'® however, at an early date impugned this belief
for he found that the mother liquor of a strong copper sulphate ammonia
wash caused no injiuy to the grape vine, the strawberry and "various
other vegetables." A glance at table 18 will show that ammonium sul-
phate can not possibly be the cause of the toxicity of the cuprammoniums
in which it occurs, since the amoimt of ammoniiun sulphate formed could
not exceed 0.53 gram for every gram of cupric sulphate employed. But
if ammonium sulphate can not be the cause of the toxicity of the copper
sulphate and ammonia wash or Johnson's mixture, the data presented
in the table show that it possesses a peculiarity to which we may well
devote a moment's attention.
" Pearson, A. N. In Report on the experiments made in 1888 in the treatment
of the downy mildew and blackrot of the grape vine. U. S. Dept. Agr., Sec. Veg.
Path. Bui. 10: 18. 1889.
1917]
BcTLEs: The Cupiluiuonidh Washes
BS*et 0/ flew a*d quick ttryitin on tht loneily
per cent of injury
$vtpfuiU, erpratttd tn
■•
euin
™>,«n.o,^„.o-
4p««Dt
Iperont
lp««t
0.1 p««i»
Col.«. 1
B..„ /
Quickly
Slowly
Quirkly
Slowly
Quickly
Slowly
91
24
63
25
73
SO
19
28
4
34
22
24
0
16
1
8
6
0
0
0
TooMto
{
0
0
Bfftett on tomato, CoUvt and btan of the mIowIj/ dried euprawimonimiu and their eom-
ponenli <U the $lr^nftht at trhich Ihey occur in the ictuhei, txprtiud in per cent of
256 Phytopathology (Vol. 7
Amnionium tmlphute is more toxic when drieci quickly than whrn
drietl slowly, beha\ing in this respect exiietly the reverse of the other
components of the cuprammoniums all of which are much more toxic
when dried slowly. The injur>' pro<luce<l by quick dr>ing may \x* vvry
markcKl and the illustration, plate VII, fig. 2, will add but emphasis to thf
striking figures in the table. The explanation of this peculiarity is that
ammonium sulphate is slightly hygroscopic and when dried quickly with-
draws water from the leaf which vnlt» and dies, but that when drietl
slowly equilibrium lx»tween the c(»lls of the e])id(»rmis and the solution is
r<»ache<l by the time dr\'ing begia**.
The plant most scTiously injunnl by quick (lr>iug is the (\>leus, then
follows the lM»an, the tomato Ixing but littl<» more* seriously afTert^l by
quick dr>'ing than by slow dr>'ing, the data for 1 per cent solutions beinK
in this respect particularly ia»«tnictive. It would ap|M»ar then»fore that
the cutirl(»sof the Coleusand b(»an are not reacHly iKTineatiMl by ainni«>-
niuin sulphate while that of the tomato is easily iM'netnite<l which farts
are of cdnsidrrable inten'st since slow drj'ing which pVoiM»rly nu^asuroA
the toxicity of the salt shows that the most readily iH*netrat<Hl» leaf is
also the most siiscei)tible.
Having conchuled our study of th(; com])onents of the cuprammoniums
we are in a [)ositiou to determine the nature of their role in the toxicity
of tht»s4' washes. In order to simplify our stu<ly I havt» placinl en regard
in tabh» 19 the iMTcentage injuries pro<luc(Hl on the one hand by the
M»Vfral cupranunoniums and <»n th<* other the injur>' that wouhi have
followe<l the us<* of the com]>on<nts at the corresiMinding strengths at
which they <M*cur in the washes, the* data being in all cases for slow ilr>'ing.
The data prrM-nttnl in the tnblr confirm in large measure the \ifw of
Millardet that solubU* copp<T is the cauw* of the injur>' pnHluce<i by the
cupranmioiiiums though it would also apix*ar that anunonia lH*c«)mes
toxir when its con(*entration excetKls an amoimt thiit may or may not \^
consiiliTably in exci'ss of that normally tolerati^l by the plant amcirniil.
For instance in the cas<' of the Coleus, malachiti'-ammonium carlK»n:it«»
containing ().2S7 imt cent copper, is more toxic than the i^quivalt-nt
strength of S4)lubli' copfHT or ammonia whence it must Im* c<mclu<h*tl that
the injury is due to coinbim^l action of thr two; and it would appear
fn»ni thr behavior of this wash at other strengths that the pn's«nri' of
aimu<»nia has in all ras<*s even when its<'lf not apparently tiixic. increas«il
the toxicity <;f the copi>er. The malachit4'-amm(mium carUinate wash
aUo ^hn^l*< evidi nr«' of an a<lditive effect except at the hiwest concentra-
tion U'^mI. (Ml the other hand. Johns<m*s mixture shows that the t4>xicity
of snlubl* eiipiM f i^ re<liired by it< presence despite the fact that the c<»n-
centration of aniiii« nia is verv nearly the sjime as in malachite-amnii>-
1917] ^ Butler: The Cuprammonium Washes 257
nium carbonate; but in the case of the copper sulphate and ammonia
wash the evidence is conflicting. When we come to consider the tomato
we find that the presence of ammonia reduces the toxicity of soluble
copper in the case of Johnson's mixture and the copper sulphate and am-
monia wash, but that in the malachite-ammonia wash its toxicity is
increased except at the lowest concentration used. In the case of mala-
chite-ammonium carbonate we have the toxicity of soluble copper in-
creased at the highest concentration (0.287 per cent copper) not changed
at 0.143 per cent copper and 0.035 per cent copper and reduced at 0.071
per cent copper.
Finally in the case of the bean we find that the presence of ammonia
has in all cases reduced the toxicity of soluble copper.
The evidence is therefore in favor of the view that the presence of am-
monia isL beneficial and not injurious as too commonly supposed though
it should be noted that while this beneficent action of ammonia is suffi-
ciently general to be considered established and is independent of the
rate at which the cuprammoniums decompose, the data nevertheless
clearly show that cuprammonium sulphate, the most unstable of the
washes, is also, all things considered, less inj\u*ious than cuprammonium
hydrate and carbonates which are very stable. Soluble copper must
therefore be the major cause of the toxicity of the cuprammoniums in
the interim between application and desiccation.
B. Effect of the cuprammoniums after they have dried upon the plant and
are subject to the action of the weather
After the cuprammoniums have dried upon the plant there will be
present on the leaf only a copper carbonate or hydrate in the case of the
washes prepared from malachite, but in the case of those prepared with
cupric sulphate (copper sulphate and ammonia wash, Johnson's mixture)
ammonium sulphate not in excess of one-half of the cupric sulphate
employed will also be present, and in the case of the washes prepared
from Burgundy mixture we will have besides the copper salt both sodium
bicarbonate and sodiiun sulphate present, the former to the extent of
0.41 gram for every gram of cupric sulphate taken, the latter to the ex-
tent of 0.56 gram for every gram of cupric sulphate used.^^ In two
instances, therefore, the dissolution of the copper in meteoric waters is
not affected by the presence of a foreign substance, in two instances the
foreign substances (sodium sulphate and bicarbonate) are neither toxic
** The calculations are based on the formula (vide Bedford, Duke of, and Pick-
ering, S. U. loc. cit., p. 86). 5 CUSO4 5 H2O+8 NajCO,. + 10 HjO = 2 CuCO,, 3
Cu(OH),-f 6 NaHCO,-f 5 NajSO* + 105 H,0.
268 Phytopathology [Vol. 7
at the strengths at which they are found in the washes or solvents of the
copper salt, and in two instances the action of meteoric waters is heightened
by the presence of ammonium sulphate which is a solvent of inflolul>le
copper salts but non-toxic itself at the strength at which it occurs in the
washes. It is, therefore, clear that the toxicity of the cuprammonium
washes after they have dried upon the leaf can only be due to the prei^-
ence of soluble copper. Whence it will only be necessary, in order to
determine the magnitude of this toxicity, to ascertain the dogrec of
solubility of the copper in the several cuprammoniums as compared with
that of a soluble copper salt, and, as already indicated, cupric sulphate
is the only conunon inorganic salt of copper that satisfies the condition!<i
under the experimental methods employed. It is desirable in teeing
the relative solubility of cupric sulphate and of the copper salt of the
dried cuprammoniums that an organism sensitive to soluble copper be
employed as indicator. The conidia of Plasmopara viiicola are, as \»
well known, extremely sensitive to soluble copper*' and are admirably
adapted to thin purpose and were employed in obtaining the data presented
in table 2(). The data were obtained by spraying microscopic slides dSUsr
the manner tlcscrilN'd by Ke<ldick and Wallace'' with the solutions to be
testeil, allowing them to dr>' spontaneously at room temperature and
putting in Her\'ice not earlier than twenty-four hours after the fungicides
had dricMl. In making a test the spores were washcni with distilled water
into a l)eaker from leavers just freshly gathcre<l. Small drops of water
with spores in sa^^pt^nsion were taken from the l)eaker, and place<i on the
slides which were then incubatiHl at or near the optimum temperature
for iiidirei*t geniiination.'^ Simultaneouf^Iy a witness was always pre-
pare<i so that the vitiility of the spores could lx» properly judged and no
exiM^riment was considere<l in which the germination in the witness prove<i
low. Finally it should l>e noted that the data given in table 20 are the
mean of five ex|M*riments except in the case of the malachite-ammonia
wjish with which only two testn wore made.
Tlu; d:ita show that the presence of anunonium sulphate increases the
toxicity of a wiu^h. the unit copper in copfXT sulphate and ammonium
carlM)i)ate U^ing mon; toxic than the unit coppcT in malachito-ammonium
^ .MillnrtJrt, .\. niid (iiiyon, l'. Traitniifnt du mildiou par Ic m^lAnge dr vulfAtr
d<» niivrr <»t dt» rhnux. Jourii. agr. prut. 2: 7<X». 1SS.5.
Wtithrirh. K. IVUt die KinwirkiinK von MotnllHaltcn und Siuren auf die
KtMinfahigki'it der Sixtrrn einifcrr don ViTbrritetiitcn pjiraiiitiirhcn Filw unarrer
Kultun>flnn£4>n. //<*it. rnaiiienkr. 2: ir>-:U. Hl-IM. l.Hirj.
" Hrddirk. I>. and Wulinn*. K. On a InUiratory method of drtermtninK the
funKit'idal valui* t»f a ^pr:ly niixturr or nolution. S<'ienre n. s. SI: 798. 1910.
** MflliiiH. I. K. (ii'nninatinn and infrctinn with the funguf of the latr blifcfat
of p«>tato. WiHconmn Agr. Kxp. Sta. Tech. liul. S7: 'M\. ltU5.
1917]
Butler: Thb Cuprammonium Washes
259
carbonate. It will be also noticed that the highest toxic value for the
unit copper occurs in the copper sulphate-ammonia wash, though on
account of the presence of ammoniimi sulphate it is impossible to tell
to what extent this high value is due to the presence of this solvent,
to what -extent to the basic cupric sulphate. Judging from the increased
toxicity of cupric carbonate in the presence of ammoniiun sulphate it
would seem that the high toxic value of the unit copper in basic cupric
sulphate is due to the presence of ammonium sulphate. However this
may be, the table shows conclusively that the unit copper in the cupram-
monium carbonate washes has a lower toxic value than in a cuprammonimn
hydrate or sulphate" wash.
TABLE ao
Relative toxicity of the cuprammonium washes and cupric sulphcUe to the spores oj
Plasmopara viticola
rUirOICIDB USED
Malachite-ammonia
Malachite-ammonium carbonate
Copper sulphate-ammonia
Copper sulphate-ammonium carbonate
Cupric sulphate
LETHAL STRENGTH
per cent Cu
0.0033
0.0057
0.0027
0.0035
0.0019
SOLUBILITT or
COPPER
Relative nutnbera
58.13
33.33
70.42
54.34
100.00
Knowing the toxicity of cupric sulphate to any given higher plant
it may be calculated readily from the data given in table 21 at what
strength the cupranmioniums would have to be used in order to prove
uninjurious after they had dried on the fbliage and are wetted by meteoric
water. Taking the plants used in the experiments with cupric sulphate' •
we arrive at the results shown in table 21, the data for the cupric sul-
phate being obtained from table 15, either directly or by extrapolation.
The data presented in the table show the strength at which the several
cuprammonimns could be applied to foliage without producing injury
after diying due to dissolution of the copper in meteoric waters. The
question that we must now consider is whether these strengths are supe-
rior to or smaller than those at which injury occurs between time of
application and time of drying. If inferior then the maximum strength
at which the cuprammonimns can be used will be governed by the sol-
ubility of the copper after drying; if superior, then the degree of injury
•* There is no reason for supposing that modified eau c61este or Burgundy mix-
ture-ammonium carbonate would possess a toxicity materially different from their
prototypes mentioned in the table.
» See p. 251.
260
Phytopathology
[Vol. 7
produretl while dr>nng will limit the strength at which theme funfdcidcs
niay be useil safely. The data presented in table 22 will permit us to
answer the question raised.
The data show that in the case of all four fuiigicides the degree of tol-
eran(*e of the drie<l fungicides must limit the strength at which thc>' cao
be aseil.
TABLK 21
Calculated tolerance of jAatitn to dried cupramfnoniumn
ri.JiXT CMPU>TED
j crPMic
nrLPHATC
UHIED
MUJWLT
M)N-TOXir
MAXIMTM STRCNOTH AT WHICH rrraAlllt«>Vtt Hi
COt'LO BB IHBD
I
Malarhite- '
nmniunia
Malachitt«-
aninioniuni
carlx>iiat«' i
C'«»pprr
Hil|»h»l«>-
ammonia
•ulpnal*^
ainn>««n«uM
rartmojai*
CflllMlS
T(»iiiat<)
CaiiliflowiT
PrlarKoniuni
OxaliM . .
prr cent Cu p*r r^nt Cu
I 0 (K)?.") 0 0110
[ 0 (M)?.') 0 (HID
0 (NN)7 0 (N)12()
0 (KM 7 0 ().>».■)
0 OOP.) ! 0 (KXTJ
0 0070 i) ()\:i'}S
fur ernt f^u
0 (n?2.*>
0 02-r)
0 (xrji
0 int'ii
0 (K):i7
0 (n.>:{7
\ptr rent Cu
- 0 oKm
I 0 01<N>
: 0 (XNr.lO
0 (urio
; 0 00-3)
i 0 0112
Mr rr<»f *'%
0 OI.'iH
o oi:{H
0 IHMJH
o a\\:i
0 miM
o oi4:>
TABI.i: 22
Affiouut of cop lie r toUratrd in thr Atr*Tol ruprammoni urns during and after drv^t^g
( OLM n
TDMATO
• ft«^
rcNdu mr. i ii>.d
(%>p{)«*r tokratril j (\>p|irr |o|rratr«l i ('4*|>prr to|rrat«»l
Dunne
dryins
4ir>ib<
' jtrr rrut jt^r frnt ■ per rent • ptt rrnl
Cu ru Cu Cu
.\!al:irhit<'-aiiiiiionia
r rrnt
Cu
, 0 01>«W 0 Oi:i.">.S: 0 IKTJ7 ■ 0 Ol.'USi 0 02S7 O <«t<J
.Malarhitr-aiiitnoiiiiiiii rarlM>ri-;
at(*
' 0 o:{7:i 0 02:17 ' o ojs7 ' o oxiT \ o oim3 n tiii:»7
(*i>p|>4-r Miilphatc-ainnionia.
0 inr.N • 0 0112 ' 0 ofki.') i o 0112 1 0 01.'>2 1 0 oirjn
C'np|MT ^ulI>hatf*-aIllInolliuI^ ' I I
rarlMiiiatf i 0 02.Vt 0 OU.'i 0 OtWio
I I
0 0145 : O (r2*> U UUI
(\ Injurtj hij niiihtivr cfftct
A.*« wr \\:\\'r h*4*ii th(' ru|>r:iiiiinoiuuiiis may pnMluce injur>' cluniig
ilryiiiju ami aftrr drying dur to di>M)hition of the cMipper in nwtoorio
^:it«T>. Whni tin* injuria pro<lur<Ml l)y thrs4» sourw^s is sufficiently
distant, thrir mutual rfTrrt> rnnaiii diMiiirt, but when thev wcur ncarlv
^inlultaIl('4>tl^ly an injury gn^atrr than that <lut» to the sum of the eflft-rt*
1917]
Butler: The Cuprammonium Washes
261
of the injury produced during drying and after drying is to be anticipated
since as SchandeH^ and Barker and Gimingham^^ have pointed out
soluble copper is more injurious to recently than to remotely injured
leaves. The data presented in table 22 show clearly that if injury is
produced during diying, injury will also follow wetting with meteoric
water. The converse is not, however, necessarily true, since the rate
of drying markedly affects, as we have seen, the degree of tolerance of
copper.
III. PRACTICAL considerations
If we consider the strength at which the cuprammoniums sffe or have
been used in practice, we will find that these fimgicides are, or have been,
as will be seen from table 23, applied at a strength in copper ranging
from 7.2 to 47 times the lethal concentration for Plasmopara viticola,
table 23
Strength at which the cuprammoniums have been most commonly employed in practice
and lethal concentration of the same for Plasmopara viticola
a
B
A
PUNOICIDE USED
Strength at
which applied
Lethal
concentration
RATIO =
\lalachitc-aininonia
per cent Cu
0 . 0493
0.0264
0 . 0270
0 0254
per cent Cu
0.0033
0.0057
0.0027
0.0035
1: 14.9
Malachite-ammonium carbonate
CoDDer sulohate-ammonia
1:4.63
1:47.03
Copper sulphate-ammonium carbonate
1:7.25
Now since the cuprammoniums may all be considered equally adhesive,
as regards resistance to mechanical shock, because they form precipitates
composed of particles of nearly like size, the concentration at which they
are applied should bear a definite relation either to their efficiency or
efifectiveness.'* But an inspection of table 23 will immediately show that
no relation exists between lethal concentration and strength of applica-
tion whether we take as our criterion efficiency or effectiveness. The
" Schander, R. Uber die physiologische Wirkung der Kupfervitriol Kalkbriihe.
Landw. Jahrb. 33: — . 1904.
•• Barker, B. T. P. and Gimingham, C. T. The action of Bordeaux mixture on
plants. Ann. Appl. Biology 1: II et seq. 1914.
*• The efficiency of a fungicide depends both on the solubility of the copper and
its toxicity when in solution. Usually, though not necessarily always, solubility
is a measure of toxicity and conversely. Effectiveness depends on the power of a
fungicide to withstand weathering, that is, to possess adhesive properties, while
at the same time yielding sufficient soluble copper to give protection from a specific
organism or organisms.
262
Phytopathology
(Vol. 7
formulae in use should therefore be amended so as to give a numerical
relation between lethal concentration an<l strength of application.
liCt UH first of all determine the factor required to give maximum
effectiveness.
Since we have been unable to assign a value to this factor frr>m the
data presented in table 24, it will be necessary for us to determine it in-
directly and this we can do from our knowledge of Bordeaux mixture.
Acconling to a recent French enquiry*** a 2 per cent Bordeaux mixture is
ncccssar>' to give adequate protection from Phmnopara viticola in yca»
favorable to the development of this parasite though in years when in-
festation is not severe 1 per cent mixtures meet more or less satisfactorily
the re<|uircmcnts of practice. In the United States, on the other hand,
both Plasmopara viticola and Phytophthora infeatans may be .satisfac-
torily held in check by 1 per cent Bortleaux mixtures though 1.2o per
cent mixtures are ako commonly emplc)y<Hl. We may therefore in all
propriety take a 1 per cent Bordeaux mixture for our standanl of
comparison.
Now Bordeaux mixture 1: 1 is toxic to Ixjth Plasmopara viticola and
Phytophthora infiHtans*^ at 0.0039 per cent copper which gives us a factor of
64 as the re<juir(»ment of practict^ for adcKiuaU^ protection. Accepting this
factor of M for Bonicaux mixture the factors necessary to apply to the
cupramnumiums, due regard Innng taken of the n^lative (efficiency of the
unit coppcT, in ordiT to obtain satisfactory' pn)tection would then be as
indicato<i in table 24.
TABI.K 21
Fariarn by which the lethal nmrtntnitiorm oj the aereral cufframmoniumM mu*! he mui-
tipliiii in onier to obtain adequatt ftrotertion
rt-KUk'iDC rtiKD
UCTNAL
COMt KWTIIATIOM
lionli'AUx uiixtun* 1:1
Miilnrhitr-iiinniunia
Mnlarhit4*-Hiiiiii«)iiiuiii rarU«imtr
( *t>p|M*r Milphiitc-iiiiiiiiotiiti
C'oplMT Milphatc'-aiiiiiuiiiiiiiii rnrlNiiiatt'
0 n(»9
0 (X»33
0 ortt7
0 0027
0 0035
r4rToa
64
53 7
93 4
44 I
56 9
Fntiii th«' ilata giv(*n in table 24 one (*an n^adily d(*teniune the stn*ngth
at uhirh the riipraiiunoniuiiis hhould Im* us<*<1 in pradice in order to ul>>
tiiiii :i pnae<'tion ^*ul^^<tanti^llly (H{uivalent to that given by Bonicaux
iiiivtun'. :tn<i when the data m» obt^iined :u'e eoiiipanNl i^nth the strengths
* (':ipiiM. J. Ia'a traitiMiieiitH ilu iiuMiou. Urv. d<* vit. 44:302. 1916.
•' WiM-niiHin Agr. i:xp. Sta. IVch. Hul. 37: M). 1915.
1917]
Butler: The Cuprammonitjm Washes
263
employed in practice, as in the following table, we find that the calcu-
lated strengths are from fom* to twenty times greater than those that
have been actually used except in the case of the copper sulphate and
ammonia wash when the agreement is close. But since the cuprammo-
niums have not afforded, at the strengths used in practice, protection
commensurate with Bordeaux mixture and we are now in a position to
understand why they have not, the formulae employed should be emended
so as to approach the calculated values indicated. But the actual
strengths employed will be determined by the tolerance of the plant
sprayed to the fungicide in the interim between appUcation and desicca-
tion, and to the solubiUty of the dried wash in meteoric waters. We will
first of all, consider the latter case.
The dried cupranunoniums must not, of course, yield on being wetted
more soluble copper than^the sprayed plant will stand. Let us accept
table 25
Strengths at which the cuprammoniums are used in practice and strengths at which
they should he employed in order to give protection equivalent to Bordeaux mixture
rUNOlCTDK USED
BTRENQTH USED
IN PRACTICE
STRENOTH EQUIVA-
LENT TO 1 PER
CENT BORDEAUX
MIXTURE
Malachite-ammonia
per cent Cu
0.0493
0.0264
0.1270
0.0254
per cent Cu
0.1772
Malachite-ammonium carbonate
0.5323
Copper sulphate-ammonia
0.1190
Copper sulphate-ammonium carbonate
0.1991
for the sake of concreteness, the value for the tomato 0.0075 per cent
copper. This percentage of metallic copper is yielded by a 0.045 per
cent copper sulphate and ammonia wash under laboratory conditions
but the tolerance of the tomato under field conditions may be safely
placed at 0.125 per cent, since rains even of moderate intensity will carry
away appreciable amoimts of the fungicide. And since in the other
cuprammoniums, as we have seen, the copper is less soluble than in the
copper sulphate and ammonia wash, we may safely use them at the
same relative concentration which would then give us the following as the
permissible strengths at which they can be applied without injury from
soluble copper resulting to a plant tolerating 0.0075 per cent soluble
copper. As will appear from a consideration of table 26, the calculated
values for soluble copper tolerated are lower than those obtaining in
practice for the malachite-anamonia and copper sulphate-ammonia washes
and higher in the case of the two cuprammonium carbonate washes.
2G4
Phytopathology
[Vol. 7
The (lata presented in table 26 further show that it is impotwible to
apply the ouprainnioniums at strenjijth*s equivalent in effectivencsw to 1
per cent Bordeaux mixture when the plants spraytni will not tolerate niorp
than 0.(K)75 per cent soluble copper since in onler to obtain equivalence
the plant sprayed would have to tolerate 0.0158 per cent soluble coppiT.
We have adniitteil that, under the conditions of practice plants will
tolerate cupraninioniunis 2.8 times stronger than tolerated under critical
conditions. It remains now to be seen whether cuprammoniumA <»f this
strenjrth can l)e applied without injury resulting during dr>nng. The
data present(Hl in table 27 show that in the cjise of the tomato uxi<ler
the conditions of quick drying, all the cupranunoniums except tlic mala-
chitt^junmonia wa.^h can Ix* us(»d at tiie nH|uired concentration, hut that
under the conditions of slow drying inalachite-ammonia and malachite-
anunonium carbonate, the latter particularly,* are toxic at a lower con-
centration than that deinaii(lcd. In tlu^ case of the Coleus we find <|uick
TAIU.K 2«
Strt'figthM at which tht rupramtruniiumM may be UKcd withoui an iujurinu^ ntHi»uftt •»/
mtluhlr rop/HT forming on wetting with meteoric water, a plant rrnixtant Itt ft mfTS
f}er cent foluhle eop/nr in ing presupjHtsed
r\ sou tut. \ nr.n
•»TH» N«.TH
TOM M«T> II
•TRrN««TN ' •• t*
Nf.'iiarliitf'-HininnTii:!
Mal:ir|iitc-;tnifii<iiiinin ('Hr)N)iiat«'
( *op|MT »>ul|>h:it4'-aiiiiii<»ni}i
i^opfMT huiphati'-aiiiiiioiiiiiin <*:ir)M>ti:it<*
/Mr rrnt f'u
0 (WIT
0 mio
I
(» ova
0 O.Y>l
drying jH-nuits the us4' in evrry rast' (»f MrongcT s^ilutions than tolenifnl
in the driisl wa.*<h, whil<» tlio n»vrrs<» is the vh-a* if .slow dr>'ing is p4*nnitte«l.
In the ca»* <»f the Imviu tin* plant will tolerate' Wronger dr>'ing than dri«tl
w:i.»*Im»s. In tho ras4» of thr tomato and ( 'oleus the wjislM-s cjin all lie u?«*«i
at 11.7 tiinrs their toxi<' <'on<M»nt ration to Plasmoimra vHicvia when «lri«*«l
(|uirkly but in tlir ra>4' of the iH'an thry can In* appli<'<l at only 3.^i tim«-!«
tln'ir K'thal «*onrrnt ration with safi^ty, no matter how .*ilowly or c|uirkly
th«' w:ls|m's are driiMJ. It siM-ins, thrn»f<»n», <'lrar that the cuprainmoniunu*
can n«»t Ik» ron.Md«*nNl a,< efT«»rtive as Hordraux mixtun* for the ctmtnJ
of pani'^itir organisiiLs whirh «io not n^juin' a <*onct»ntnition in S4»lublo
cop|M*r gn\atcr tlian that yi<»ldc.d by the latter. We have now t«i C'»n-
hidiT thf cupraminoniums fn»in the |M»int of view of efficiency.
TIm' unit ropjxT in the (*upramm<M)iunLs has generally IwH'n con.sidiTetl
iiion* rtficicnt than the unit ropfxT in Hord<'aux mixtun*. and this opinion
\^ mhloubt<H||y well grounded wh«*n thr cuprammoniunts which an* \\t:¥>
ti<*all\' neutral as scMin as dry an* (M^mpan^d with alkaline l^mleaux mix-
1917]
Butler: The CuPRABfMONiUM Washes
265
tures in which the copper is without action during the time required to
reach neutrality; but when compared with neutralized or neutral Bor-
deaux mixture at the lethal strengths to Plasmapara viticola the difference
in favor of the cuprammoniums is indeed small. In the most efficient
wash (copper sulphate and ammonia) the unit copper has a value only
1.44 times that of Bordeaux mixture and in the least efficient (malachite-
ammonium carbonate) it is 1.46 times less active. But in fimgi resistant
to copper the unit copper in the cuprammoniums may be manifold that
of Bordeaux mixture, the highest values being given by the copper sul-
phate and ammonia wash. For instance in the case of the uredospores
of Puccinia Antirrhini the efficiency of the unit copper in the copper
sulphate-ammonia wash is very much greater not to say infinitely greater
than the unit copper in Bordeaux mixture since it would appear that the
latter is non toxic at all concentrations.^^
TABLE 27
Amount of copper tolerated in the cuprammonium washes during drying by the tomatot
Coleus and bean
TOMATO
COLBUS
BBAN
rUNQlCIOB USBD
strencth of dried
wagb tolerated
StreoKth
tolerated
Strencth of dried
wash tolerated
Strength
tolerated
Strencth of dried
wash tolerated
Strength
tolerated
Wash dried
quickly
Wash dried
slowly
Wash dried
. quickly
Wash dried
slowly
Wash dried
quickly
Wash dried
slowly
Malachite-ammonia. . . .
Malachite-ammonium
carbonate
per e^nt
Cu
0.0386
0.0668
0.0317
0.0410
per cent
Cu
0.0359
0.1437
0.0635
0.127
per cent
Cu
0.0327
0.0287
0.0635
0.0635
per cent
Cu
0.0386
0.0668
0.0317
0.0410
per cent
Cu
0.0718
0.1437
0.0635
0.0635
per cent
Cu
0.0298
0.0315
0.0228
0.0254
per cent
Cu
0.0089
0.0154
0.0073
0.0101
per cent
Cu
0.0301
0.0229
0.0203
0.0279
per cent
Cu
0.0287
0 0183
Copper sulphate-am-
monia
0.0152
Copper sulphate-am-
monium carbonate. . .
0.0228
It is therefore clear that in the control of parasitic endophytes the
spores of which are highly resistant to soluble copper, the cuprammoniums
may be of very considerable value provided the plant to be sprayed will
withstand the concentration demanded. But except in those cases where
Bordeaux mixture is non toxic to the parasite from which protection is
sou|2:ht it does not seem to me that the cuprammoniums as a class or any
one cuprammonimn in particular possess merits sufficient to warrant
** Doran^ W. L. Controlling snapdragon rust. Value of copper and sulphur.
Florists' Exchange 48: 501. 1917.
266 Phttopatholoot [Vol. 7
their emfdoyment in practice, especially ainoe the quality of inconspio*
uouaneeB can be obtained quite readily with Bordeaux mixtures 1 : alkalinity
Bordeaux mixture 1 : alkalinity is no more conspicuous than the copper
sulphate and ammonia wash when the same amount of copper is applied
per square meter in both cases but since the latter usually wets the foliage
better than the former it appears to be less highly colored, due to the
fact that the copper is spread over a larger surface. Whenever Bordeaux
mixture 1 : alkalinity does not wet the foliage sufficiently to offer the proper
degree of inconspicuousness, the wetting power oi the mixture may be
increased by the addition of an infusion of quillaia (i.e., saponin) or a
small amount of casein. It is therefore not necessary to resort to a cupram-
monium when an inconspicuous copper fungicide is required.
SUMMABT
1. The cuprammoniums met with in practice belong chemically in
one or the other of the following groups: (1) cuprainmonium sulphate;
(2) cuprammoniuni hydrate, and (3) cupraninumium carbonate.
2. When decomposed by drying cupranunonium sulphate deposits
the copper as a basic sulphate, cuprammonium hydrate yieldB a copper
hydrate, and cuprammonium carbonate a copper carbonate.
3. (^uprammonium sulphate is very unstable; cuprammonium hydrate
and carl)onatc very stable.
4. The cuprammoniums are more toxic when slowly than when quickly
dried.
5. The toxicity of the cuprammoniums during drying and on weather-
ing is due to soluble copper.
6. The copper sulphate and ammonia wash and Johnson's mixture are
less injurious than the malachite washes.
7. The cuprammonium washes arc more efficient and effective than
Bordeaux mixture when large amounts of soluble copper arc required to
give pn>tcction.
8. The cuprammonium washes arc less effective than Bordeaux mix-
ture when hmall amounts of soluble copper suffice to give protecticin hut
with the exception of the mala(*hite-ammonium carbonate wasli are
slightly more efficient.
9. The relative efficiency of the unit copper in the cuprammoniums is
in decreasing onier as follom-s: Copper sulphate-ammonia, malachite*
aiiunonia. copper sulphute-ainmonium curlxinate, malachite-ammonium
carbonate.
10. Tlie cuprammoniums nmy be used at 11.7 times their lethal con*
1917] Butler: The Cxtprammonium Washes 267
centration for Plasmopara viticola on plants not affected by 0.0075 per
cent soluble copper.
11. The cuprammoniums are of limited practical applicability and
should not be used' in lieu of Bordeaux mixture whenever the latter yields
sufficient soluble copper to give protection.
New Hampshire Agricultural Experiment Station
Durham, New Hampshire
BIBLIOGRAPHY
AuDOTNAUD. Le mildiou et les composes cupriques. Prog. agr. et vit. 1885.
Bedford, Duke of, and Pickering, S. U. Woburn Experimental Fruit Farm
Report, 11: 1-1429. 1910.
BouRCART, E. Les maladies des plantes, 372-382. 1910.
Chester, F. D. The copper fungicides, Joum. Myc. 6: 22-24. 1891.
Gastine, G. Emploi du carbonate ammoniacal de cuivre contre le peronospora.
Prog. agr. et vit. 8: 114^117. 1887.
HoLLRUNG, M. Handbuch des chemischen mittel gegen Pflanzenkrankheiten.
1898.
Johnson, S. W. Note by the Director. Connecticut Agr. Exp. Sta. Rept. 1890: 118.
1891.
LoDEMAN, E. G. The spraying of plants. 1899.
MoissoN, H. Traits de chimie Min^rale, 5. l905.
Patrigeon, G. Nouveaux proc6d^ de traitement du . mildiou. Joum. agr.
prat. 1: 881-882. 1887.
Penny, C. L. The preparation of ammoniacal solution of copper carbonate. Dela-
ware Agr. Exp. Sta. Bui. 22. 1893.
Pickering, S. U. Note on an anmionio-copper sulphate. Journ. Chem. Soc.
Trans. 48: 336-339. 1883.
Pickering, S. U. On the basic sulphates of copper. Chemical News 47: 182. 1883
288 Phytopathology [Vou 7
DESCRIPTION OF PLATES
Plate III
Effect of the malachite-ammonium carbonate waah containing 0.28 per oeat
Cu on the tomato var. Bonny best. Plants on the left dried quickly, planta on
the right dried slowly. Photograph taken twenty-four hours after tlie fungicide
was applied.
Plate IV
Kffect of the malachite-ammonia wash containing 0.28 per cent Cu on tlie lomato
Tar. Bonny l>ost. Plants on the left dried quickly, plants on the right dried tlowlj.
Photograph taken twenty-four hours after wash was applied.
Plate V
Tomato var. lionny liest photographed twenty-four hours after being spr»yed
with 1 per cent cuprir Mulphate. Plants on the left dried quickly, plants oo the
right dried slowly. The plants dnod quickly showed slight scorching of ytning
leaflets at the time of making thf photograph but the injury was not sufficiently
marked to show in the plate; the plaiitrt dried nlowly were, on the other hand, very
seriously injurc<l all the leaves UMng with(*rt>d and flaccid.
Plate VI
Kffect of ammonium hydn>xid containing 4.11 \H»r cent ammonia on the tomato
Tar. Bonny best. Plants showing no apparent injury dried quickly, withered plants
dried slowly. Hiotograph taken twrnty-four hnurs after treatment.
Platk VII
Fio. 1. Kffect of ammonium rarUinate containing 0.96 per cent ammonia on the
Coleus var. (iolden l)edder. Plant on tin* right dried quickly, plant on the left
dried slowly. The photograph wan taken furtyHMght hours after the salt was
sprayi*d on the plants.
Fig '2. Kffect of a '2 |mt c«*nt Holution of aninionium sulphate on the bean var
Dwarf hortiniltural. Plant on the left dried slowly, plant on the right dried
quickly. Photograph taken forty^Mght hours after treatment.
Pi.\tk VIII
<iraphH t*ho\%iiiK the relative toxicity of f|iiickly iitid mIohIv dri«*<l solutions of
cuprtr trilph.'ite to the < 'oleiiH. tiHiiato. Ox.mIih. Perl:irKoiiiiiiii. lM»an and cauliflower
Pl.VM I\
(fr:ipli!« -tliiiuiriK rrLitive toxicity of i)iiickly and sIohIv dried m^hitions of amm«>-
niuiit li\<lrii\iii :irid aniinoriiuiii carl)on:ite to the )N*:iii and cuulitlowrr.
Pi.\Ti: \
(iraph- »lif»uiiik; ril.itive tn\irir\ nf i|tiii-kl\ .iihl •tliiulv 1 1 ned mil utii>ns of ammo-
nium h\ilri>\til titil :tiiitiioiiiiiiii i\irlH>ii.iti' lti ttio tnni:it«t :iitd <^>Ieus
rtlYTOPATIIOIXX>Y. VI t
FHVTOPATHOIJJGV, VII
PUATK VIII
HtUOTYM 00. tOiTO*!
^ASHKS*
PHYTOPATHOLOGY, VII
PLATB X
O. B. d«l.
BuTUER : Th« Cuprammonium Washes
HtUOTVPt 00. MtTOM
1917] Boncquet: Bacillus morulans 271
from diseased plants or portions of plants, it has not been possible to
produce curly top by inoculation with cultures of this organism; that the
same organism has been isolated from the surface of beet seed, the sur-
face of normal sugar beet leaves and from the soil about the roots of
sugar beets. Also that certain bodies which seem to represent the same
organism have been found in great abimdance in curly top sugar beets,
in the interior of the sieve tubes, accompanying a specific lesion in the
phloem, and that similar bodies, in varying but much less abundance, were
found in the same tissue in supposedly normal beets or those with various
morphological irregularities of the foliage (fig. 1). Whatever may be the
entire significance of the organism in question, its peculiarly abundant
occurrence in connection with the sugar beet and its apparent relation to
curly top have seemed to justify its careful study, and it is the piupose of
the present article to describe more in detail the characteristics of this
species, to which the i^une Bacillus monUans has been given.''^^- ..'
THE DISEASE
ft
The disease of sugar beets called curly top is of annual occurrence
throughout the sugar beet growing regions of Colorado, Utah, Idaho and
California. The severity of the disease, however, varies greatly from
year to year. Some years veritable disasters are produced by curly top,
thousands of acres of sugar beets being totally destroyed after all the ex-
pense of preparing the ground and planting the crop has been imdergone.
Symptoms on leaves
The comparative size of the inner and outer leaves is altered. The in-
ner leaves are dwarfed, the petiole especially becoming shorter and flatter
than the normal, while the outer leaves, if already full grown before the
disease becomes apparent; maintain their natural size and shape and, for
some time at least, their color, although they may finally turn yellow
and die prematurely. The first symptom of abnormality plainly visible to
the eye is a distinct transparency of the finest venations of the youngest
leaves^ This transparency starts at the base of the leaf blade. Gradu-
ally the abnormality works higher on the leaf until finally the whole leaf
is aflfected. The youngest leaves are first to suffer; the older ones (such
as are not already full grown) show the symptoms as their expansion and
growth progress. Almost simultaneously with the appearance of the
transparency of the veins small warty protuberances appear upon the
veins on the under surface of the leaves, eventually even upon those which
are of the smallest size visible to the eye (fig. 2) . The margins of the af-
fected leaves then begin to curl slightly upward so as to expose the lower
1917] Boncquet: Bacillus morulans 273
several abnormal bendings; the angle of each bend is slightly swollen and
if the rootlet is far advanced the swollen region appears to be necrotic
(fig. 3). When the main root is cut transversely the successive rings of
vascular tissue appear discolored. On careful examination it will be
observed that the phloem is the only part of the vascular system which
suffers severely. This phloem discoloration is observed more or less
throughout the whole system in the veins as well as in the roots. This,
however, only becomes apparent to the naked eye when the disease has
reached its severest aspect.
Cause of the disease
It was E. D. Ball* who discovered that the sting of the insect EuteUix
tenella Baker is a necessary factor in the causation of this disease. His
observations were confirmed by Shaw,' and very fully tested and confirmed*
by Smith and Boncquet. The latter, however, together with Hartung,*
proved a fact which had previously been suspected, that EuteUix teneUa
is not the fundamental factor in the causation of this disease, but rather
must be a carrier of a second factor, presumably a parasitic micro-organ*
ism. This discovery made very important a thorough search for and study
of all micro-organisms which possibly could be found in connection with
the disease, and it is with this portion of the study of curly leaf that the
present article hais to do.
BACTERIOLOGICAL INVESTIGATIONS
The methods and detailed results through which the conclusion was
reached by the writer that Bacillus morulans inhabits constantly and
specifically plants affected with curly top, as well as occurring in certain
other situations may first be described. It was decided at the outset to
make a very thorough and accurate search for any organism which might
be present in the tissues of plants affected with the disease. The unfav-
orable or at best imcertain results reported by previous investigators
along this line led to the belief that the problem would be a diflScult one,
* Ball, E. D. The beet leaf hopper. Utah Agr. Exp. Sta. Ann. Rept. 16: 16.
1905.
The Genus Eutettix. Proc. Davenport Acad. Sci., 12: 41 and 84. 1907.
The leaf hoppers of the sugar beet and their relation to the "curly leaf"
conditions. U. S. Dept. Agr. Bur. Ent. Bui. 66, pt. 4. 1909.
» Shaw, H. B. The curly top of beets. U. S. Dept. Agr. Bur. PI. Ind. Bui. 181.
1910.
* Boncquet, P. A. and Hartung, W. J. The comparative effect upon sugar beets
of Eutettix tenella Baker from wild plants and from curly leaf beets. Phytopath»
h\ 348-349. 1915.
274 Phytopathology (Vol. 7
but at the siiine time a Rtudy of the nature of the disease had led to a wry
strong feeling that some parasitic micro-organism, of which the* in.*««*ct
Eutettix tenella was presumably a carrier or secondary host, muKt lie in*
volveti in this disturbance. Assuming then that the sought-for <>rganL<ni
would l)e an obscurer one and difficult to demonstrate by onlinar>' cuhund
or histological methoils, various siXMrial and somewhat elaborate culture
methods were attempte<l.
Preparation of media
The following meilia which seemed most promising fortius purpoM* wen*
preparcMJ:
FilUrni hrrt juicr. For thi» purpom* thr plniitH from which the juic«» wan (lf«irnl
'wcro wiiHhr<l nM thorouKhly uh poHsihle in Hterilisod wntor Aiid then icri>un(i fiiH* in i
ninit ^rindor. In niont ram^s 1(K) vv. of (liHtillcMi wntor wait a(ldf*<i to rarh 1(1) icmn.t
of hcrt pulp and t)i«* niasH w»h thon covered and h*ft xtandinK for two houri*. Thr
rnihhtMl material waM Hii)iHc(]Uently put into clean ch(H'iM*cloth and the juir«* Mpifrfrd
out in a prcHi*. Thf juice thus o)>tained wa8 fluhitequently dilutc^l twin* it* vuluinr
with n (t Halt sf>lution. VariouH dcf^recM of dilution have \wvi\ einployni. hn«ifvrr
from th«' original juice up to alxnit ten t<» one. either in Rait aolution <ir w.itvr In
mime cas4't< tlic solution was then titrated and hroURht to the neutral p<iint «if phfn>M-
phthalein with so<lium hydroxid. This juice was now clear, nliichtly hmvin and
passi'd easily thmufch a common filter pafn'r. After it had iN'cn filtered through
pafMT it wa> parsed throuKi) h mciliumHh'nse liiTkefeld filter camlle for pur|M»<M-» nf
sterilization. An apparatus wan e^perially devised for this purjMim*. a fitriii of whirh
IS des(-rili«>d in antither article in the presi'iit lunnlier of l*hytopath«ilofEy. In onW-r
to Ih* Mire that rill miitaminatinn had ocrurred durinK the manipulation, the tuU'«
after tillniK uen* ke|»t in the inrtitiatnr fftr two days at ^tf)''(\ It i*« Udievrd that lhi«
app:iratii!4 and method is wnrthy of eonf<ideralile employment in the preparatimk «>f
cultiin* media for use in plant patlmlni^y.
.1 *f 1*1 If, iitif.mftiihit t >/<M ^. 'rhe?«eMer4' prepared in the following manner Smnd
healthy l>eet*i Were •<i-le(ted. thoroughly eleancd and immers4*d in iNiilmg water f^r
three minute^, in \]i\^ ^ay !«terili7.inK the surface hut not heating or chanKinx in .iti\
maniiiT thr ti>stie;* dr»per in the lH«>t. They wen' thi'Ii cut into sliren mth a eafr-
full\ -'rriii/ed knif«' Kaeh ^li^e Was then put into a sterile |N*tri dtsh ini«i whirh
pri'\ i<>ii-l> lirilinaiN inittient a^ar had tn^en poured.
/iff' i> .-'.K. '1 I. Ill liuthlred KTams nf lieet leaves were cut into sm:i!l piece « :%r.<l
ImiiIi-iI fur an hmit in (I ;, htrr nf uater. W ater wai* then athh'd to make up tn I litfr
aini i< f* -' iiiilifik! !'<>r !»•■ Imiir'*. It H:i<< th«-n tiltereil thri»uith ciitton an<i .*■<■> t'«- •>!
tfii- l.i«' .\rrii* i.|i|».l til 1 litre i»f Iji'luu'?* I'ft'th. The Ijeliic's hmth haii prr% i-
«iii-l\ !'•«!. p'lfiirij in I In fiitjiiw mu: May : J icrams of I.iehiK's extract. |n g^ram* i>f
\\i'*i iMji'-M i!i'l '• k'?-ini- of Mnliinn rhloml wiTf addet] to 1 litre of matrr Thi«
riM'i.iMi ^^ •- - ii'« ii'ii ;.' \ III iiM-ih/fil tn pheiinlphthalein with iMMiiuni h\iiri>\i>l
aii'i ."•: .•:-;.•!■: ■! ■!.• 1"«' jui- • \\a«« hritiiclit up In O .'» yn^i cent acidity with
iii:i!.> I .•! I 111 - in.i fill •!: irti u ix al-n pri*p:ire<| with an increaMMl |iropiirti«>n nf
!■• « • I \*r ■■• *
1917] Boncqubt: Bacillus morulans 276
Artificial media. A protein- and peptone-free medium was composed with the
supposition that the organism did not attack the higher nitrogen compounds of the
beet. Therefore several of the amino acids were used as the nitrogen supply. Ala-
nin, leucin and tyrosin were used. Asparagin, although not an amino acid was also
considered a possible favorable source of nitrogen for the parasitic organism. All
these compounds were used in a dilution of 0.5 gram to 1 litre of water. The neces-
sary minerals were added in the following form and proportion:
Magnesiimi sulfate 0.2 gram
Ammonium phosphate 0.5 gram
Potassium nitrate 0.2 gram
Calcium hydroxid 5 cc. of a saturated solution
Ferric chlorid trace
These artificial media were sterilized in the Arnold sterilizer for fifteen minutes
upon three consecutive days. Special glycerin and glucose media were also pre-
pared. For this purpose 1 per cent glycerin was added to a part of the asparagin
medium. So also 5 per cent glucose was added to another portion. The glucose
medium was especially used for anaerobic purposes.
Other media. Ordinary media such as nutrient bouillon, potato glucose bouillon,
bean pods, milk, litmus whey, nutrient agar and nutrient gelatin were prepared ac-
cording to the standard methods.
Methods attempted for separating parasites from the plant
In order to separate the assumed parasites from the plant and obtain
them in pure culture the f ollowmg technique was used :
Surface-disinfected plant parts placed in culture medium. The usual
method employed in this sort of work consists in soaking the tissue to be
employed for a given length of time in mercuric chlorid and then washing
oflf the same with sterilized water before placing the tissue in the culture
medium. A need of standardizing this method was felt, inasmuch as
there is no assurance, as it is usually described, whether on the one hand
the disinfection was sufficiently thorough to kill all smrface organisms or
whether on the other hand the material was washed sufficiently to remove
all the mercury and prevent its being carried over into the culture medium.
The method consists in dry sterilizing a number of cotton-plugged flasks
of 50 cc. capacity or any other desired size. At the same time larger
flasks, Ukewise cotton-plugged and filled with distilled water, are made
sterile in the autoclave: Other requisites are supplies of 95 per cent
alcohol and 1 to 1000 solution of mercuric chlorid in water. The mate-
rial from which cultures are desired, after thorough wiping with cotton
swabs in 95 per cent or absolute alcohol in a photographic tray, is cut
into convenient sized fragments, but no smaller than necessary. These
are placed in one of the empty sterilized flasks and covered for a moment
with the alcohol for the purpose of removing air bubbles. The alcohol is
immediately poured off again and the flask nearly filled with mercury
276 Phytopathology (Vol. 7
•
solution 80 that all the material will be submerged. This is allowed to
remain for the desired length of time, depending upon the nature of the
tissue. The petioles and main veins of sugar beet leaves, especially fairly
old leaves, will usually stand twenty minutes, but with leaf bladen and
other more delicate material ten minutes has been found the maximum
time which can be used without too severe burning. Cut surfaces will
naturally absorb more of the solution than those protected by the natural
covering of the plant, and this can be taken into account both in con>id*
eration of the length of time which the tissue will stand without being
burned by the mercur}' and also the time neccKsary for washing it out
again. On this account it is best to cut the tissues as little as powible
before disinfecting. After the desired time has elapsed a piece of brass
wire gauze, l)eiit to form a cap over the mouth of the flask, is steriliie<l in
the flame, placeil in position and the mercur>' poured off. The flask is
then filled with sterilized water from the large flask and the water of this
first washing, after having the material well shaken up in it, is poured off
imme<iiately and more water poured in. Tlie pn>cess of pouring off and
refilling is then Continued at gradually increaseil inter>'als; the length of
each must depend ui)on the nature of the material. It was found, how-
ever, that if the amount of material in the flask is comparatively small in
proportion to its capacity, which should always l)e the case, six changes
of water, extending over a |x^rioil of two hours, is amply suflScient in every
casi\ In this case the first five changes can be made during the first hour
and the last one at the end of the second hour. If one wishes to practice
extreme caution the mouth of the flask nmy be flame<l an<i the cf>tton
stopp<T replaceil after each change of water, but this has not l)een found
ne<'i*ss:iry so long as the amount of water is sufllicient to thoroughly sub-
merg(» all th(* material. In our work the wire scret^n is usually left over
the mouth of the flask and this is freshly flanunl l)efore each change of
water. Aft4T the pnwess is completed the material is taken out of the
last watrr with flanH^i forceps, hniken into small pieces if necessar>' an<i
th<»rou^hly rnish<^i with the same instruments and dropped into the cul-
tun* licjuid.
I*i*ct culturta. It was thouglit tliat a gradual adaptation fn>ni the
plant in which tht* organism is living to the me<iium in which it was at-
tempti'd to jjmw it nii^lit I k* ntMM»ssary to insure success; therefore t he di:^
eiuird tir.Mi«- was so tran>frrnMi us to distiu^h as little as possible the cells
of tin* l>«i*t. Fnr this puri)os4» glass tuln^s were drawn out to 2 nun. dianu*-
Xvr. \\XvT >i«Tili/in>E l>y h<'at tht'V wt»n» 2iS4*iit icnlly inserted into the di*-
e:LMMi ir^ioii** of thr Ui-t to a d«*pth of 1 rm. The tul>e m'as then m*ith-
drawn. )>ririihn^ with it a |)ortion of the licM't tissue and the tenninal
part containing tlir tisMiu* \v:l<4 can^fully hniken off withsteriliied forceps
1917] Boncqubt: Bacillus morulans 277
and dropped into the medium. In this way both ends of the tissue slightly
protruded from the glass tube and came into direct contact with the cul-
ture medium. The slow diffusion of the latter was supposed to secure a
gradual change of habitat in such a way as not to hinder too severely the
growth of the parasite. Tissue was thus removed from the petiole, from
inside the root and from the larger veins of the leaf, after surface sterili-
zation with a flame or boiling water, afterwards cutting into the interior
with a steriUzed knife and then introducing the glass tube to take out a
small core of tissue.
Results of isolation experiments
The various special methods described were carried out very carefully.
The result was that in almost every instance cultures from curly top tis-
sue in all the various media described, and especially those which con-
tained glucose, showed a heavy growth after twelve hours of incubation
at 20°C. This result was practically imiform wherever diseased material
had been used. Occasionally growth also appeared in cultiu'es from sup-
posedly normal plants, but in by far the great majority of cases such cul-
tures remained sterile. This seemed to indicate that the organism was
not peculiarly difficidt to isolate, judging from its abundant growth on such
a wide variety of media. Nevertheless a painstaking work was imder-
taken in order to complete the thorough study which had been planned.
The anaerobic cultures also proved to be invaded by the same organism.
Here, however, the growth was less abimdant and extremely slow. Sev-
eral days elapsed before any colonies were visible. A great deal of effort
was further spent on work with all kinds of media but always the same
organism grew abimdantly. Contaminations naturally occurred now and
then but the fact was most decidedly apparent that the one species an-
nounced by Smith and Boncquet predominated in the tissues of curly top
plants to the practical exclusion of all others. The special culture meth-
ods described above are given in some detail, inasmuch as they may con-
tain suggestions of value in similar work. Having foimd that this organ-
ism grew so easily and abundantly upon ordinary media, the use of special
preparations was abandoned in the attempts to isolate the organism from
plant tissues and the work was carried on entirely with standard bouillon
to which 5 per cent glucose had been added. The object of the glucose was
to promote the growth of the characteristic zoogloeae of this organism,
rendering its identification in the original tubes easy without plating. By
occasional plating, as a check on the work, it was soon possible to identify
this organism very accurately by microscopic examination of tubes which
showed the characteristic ring formation at the surface of the Uquid. The
278
Phytopatholoot
(Vol, 7
TABLE 1
Results of bacterial isolation experiments by cultural methods
(Tul)eB incubated at 28H!). Disease meant curly top)
MATSKIAL
NtJiiBSK or
TVBBS
•■irut
Petioles of curly top beets
6
5 tubes developed B. wwrulans
within two days
Petioles of normal beets
6
Tubes remained clear for a w#rk.
when they were discarded
Petioles of diseased l>eets
9
8 tubes developed morulans
A leaf showing curly top symp-
5
All developed marulanM
toms on half of blade and in cor-
respondinK half of petiole.
Other side appeared normal.
This material from affected half
.
of petiole with black streaks
Correspondinfc half of blade
3
2 tul)es developed morulanM
Normal-appearinK half of petiole
6
5 tubes developed morulant two
days Inter than those from
blackened part
Norinal-appcHrinK »ide of blade
3
No development
Petiole of a diHcascd center leaf
4
3 with mofu/anir. 1 doubtful
Petioles of good-sized leaves from
10
2 tul)es frr>m 1 leaf both with morw
5 different n(»nnal-appearinf(
lann; others all clear
i>eeiM
Typiejilly diseased leaf with very
6
All developed mtrrulans
slight dark streaks in the jM'tiole
Badly affected {wtiole of name
4
All develope<l morulans
iM'et. Pieces rut out with
flamed scalfx'l
Petioles nf 4 beets from ins4'Ct-
8
Tulies from 2 plants remained
pronf cagt*. No sign of diseaM^
clear ; those f mm other 2 lieramr
slightly cloudy after m-vrral
•
days, but no mi^rulana
PftioIrK <»f 2 slightly dis<»as<'d
13
All tul>es apparently r«mtaininK
Irrivi--*. Pierrs rut (»llt with
pure cultures of mttrulana
tl.iMH'ii .Hr:tlpcl
A >fll«i\\i-h :i>t«T leaf
o
Ilemained clear
Afi «'l«l yillnwirih biM't li'af without
4
S<»me fungous growth. No morb-
•Mirly top. Tissue still s<»u!id
tans
!>•'.•» v«'«i i»f ri snrnewliat abnonnal-
0
Some growth, but no iiMiriiiij»i«
:ippi'ir:iig tn-rt but Dot With
<MirI\ top
I!i:i!*h\ - ij>pt -iriHt: !r:ivr> <»f aster,
i I.: *. » t:<7|ii iii'iiii •irihli:i. ti>iiia-
* ■ * • .»fi !•■• t ut •■ ritnl r.iddith
I't*i"!i i.t '\pi< ill\ •ii-f:iHiM| livif.
t.i- -ii -lltli ■ • ;-.!i
24 ill tubes with fungi and l>artrriA.
remainder clear. No m«>rW«in«
I founti
(i Very abundant growth of mi^^a^t
intermixf*<l with other organ usn*
1917]
Boncqxtet: BACiUiUS morulans
279
TABLE l-Continuei
IIATSRIAL
NT7MBKB Or
TUBM
BK8ULT
Petiole of slightly diseased leaf
2
Both very abundant morulanB
Blade of same between veins
2
Both remained clear
Typically diseased leaf; scraped
4
3 tubes developed morulans, 1
out interior portions of petiole
doubtful
with flamed scalpel after clean-
ing off epidermis
A young leaf visibly affected on
4
All developed morulans
one side and very slightly at
the base of the other side. Tis-
sue taken from the most dis-
eased side at base
Terminal portion of diseased side,
4
2 developed mortdans; 2 clear
less visibly affected
Slightly affected base of other side
4
1 developed morulans; 3 clear
of same leaf
Not visibly affected terminal por-
4
All remained clear
tion of last
Note.— The last four are from the leaf illustrated in Phytopathology 6: 106.
The most elaborate precautions were taken to secure perfect surface disinfection and
avoid contamination. These tubes in which gprowth appeared were plated out and
found to contain pure cultures of morulans. The leaf was perfectly sound, showing
only a slight roughening of the veins on the affected portion.
appearance to the eye of this ring, supplemented by microscopic examina-
tion, finding it to be composed of the characteristic zoogloeae, supple-
mented by occasional plating, is amply sufficient to identify this organ-
ism. A number of typical examples of isolation experiments with sugar
beets are shown in table 1.
Several himdred illustrations similar to those shown in table 1 might
be given. The results varied somewhat with the perfection of technique
TABLE 2
Bacillus morulans upon sugar beet seed
MATERIAL
NUMBBR or
TUBES
RESULT
Beet seed imported from Ger-
10
At least 7 developed an abundance
many, 1 dropped into each bouil-
of morulanSf mixed with other
lon tube, with no previous treat-
organisms
ment
Similar seed previously soaked for
10
All clear
twenty minutes in mercuric
chlorid and washed in sterilized
water
280
Phytopathology
[Vol. 7
and in individual cases, but a mass of evidence was collected to indicate
that this organism exists regularly in the interior of the foliage of migar
beets where the visible s^^mptoms of curly top occur and that it does not
develop in cultures from nomud foliage or even the normal-appearing por-
tions of partially affected leaves; also that it does not occur in the interior
of beet leaves which may be yellow or sickly from ordinary cauM«.
Cultures from seed. Many attempts similar to those shown in table
2 were made to isolate the organism from sugar beet seed. The uniform
result was that almost every unsteriUzed beet seed dropped into a tube
of lx)uilIon developed a very luxuriant growth of Bacillus fnarulans,
TABLE 3
liacillim tmnulann from noil
MATCRIAI.
NruBBR or
TCBCn
KaBn.T
Pinches of soil from alxmt the
r(H)t8 of a diseased l)eel
Pinches of soil from about the
roots of a normal U'ct in insc»ct-
pnK>f cajje
4
4
Marulana wm abundant in tpverml
of the tul)e«
Some morulan9 present in the
mixed growth resulting
Culiurenfrom soil. That the organism is present in some soils is indicated
by tho data preK»nted in table 3. The work was rather crude but cer-
tainly li. tuorulans was abundant in the soils examined.
Cultures from unsteriUzed foliage. Cultures made from unsterilised
leaves of the sugiir In^'t (table 4) show that the organism is common as a
sapro|)hyte upon the leavt^ of the* plant, but in all cases when leaves simi*
lar to these w<Te thoroughly disinfcvted no growth was obt^ned.
TABLE 4
CuUnrf* from unntfrilizrd foliage
M\ri HI«L
I.<-:ivi*«» i»f iiortiial iHTtH
Ni'MBCR or
Ti'Bm
10
saicxT
several contained an abundance
(»f moridanM
rii>t«'riliz(Hl loavt*^ of many other plants were also tried but the resulting
grtiUlh \%:i> Ml luixod that no safe rtiiirhisions (*ouhl 1m» drawn. The only
rertaiii iicvt*Inpiiu>iii< of muruUius (NTiirnMi in tnU's inoculated with pieces
of r)in>anth('iiuuii h*avi'.<.
i'ulturtt^ frttm suijtir /w«/ Imns tnth ty/Ms of disease other than curly top.
Th«- I'art that lM»ilit> n'x-iiihliiiK bacteria have b4M»n s<»en i*ith ihe micro-
iii*ii|H' in sii^jir Ui't h'av«-> not afTt-t-t^Hl with curly top, Init affected with
1917]
Boncquet: Bacillus morulans
283
TABLE 5
Cultures from sugar heel leaves with types of disease other than curly top
MATERIAL
Yellowish area of young "black
edge'* leaf taken from between
black and green portions.
Thoroughly disinfected
Similar to last, but not disinfected
Similar material, disinfected
Similar material, disinfected
Typical "mottled leaf;'' not dis-
infected. Blade, petiole and
veins
Petiole of "black edge" leaf, dis-
infected
Petioles of normal appearing leaves
from healthy plant. Very
carefully disinfected
Petiole of decidedly "mottled
leaf." Inner tissue removed
with flamed scalpel
RESULT
Both gave a strong growth of
morulans
Very vigorous and apparently
nearly pure growth of morulans
All with morulans
All with morulans
All produced morulans in abun-
dance
All appear to have pure cultures
of morulans
All tubes clear
Very vigorous growth of morulans
Further study of the organism, which was uniformly present in diseased beets
Although the organism was able to grow most abundantly on the com-
mon culture media, the peculiarity of this growth under all circumstances
was of such a nature that for some time a continual contamination was
suspected. Each separate colony seemed always to have two kinds of
bacteria, very distinct in form. Very active bacteria were always observed
at the edge of the colonies, while capsultated bacilli were generally ob-
served in the middle. Therefore, before any further study of the organ-
ism was taken up, repeated efforts were made to separate these two widely
distinct forms. For this purpose, the calcium carbonate and the India
ink method for separating the individual organisms previous to plating
them were resorted to.
Calcium carbonate. To 10 grams of calcium carbonate enough water
was added to form a milky paste. This was subsequently introduced into
a 200 cc. Erlenmeyer flask and sterilized in the autoclave. After the nec-
essary cooling several yoimg colonies of the bacteria were introduced into
the semi-liquid mass and shaken for two hours so as to separate each indi-
vidual organism from the other. From this paste, several plates were
poured in the usual manner. They were incubated at 37°C. and closely
examined as soon as any sign of development occurred.
284 Phytopatholoot [Vol. 7
India ink method. For this purpose special Chinese ink, prepare«l by
Grilbler, (Punkttusche) was used. A 15 per cent nutrient gelatin was
made and poured into clay-covered petri dishes. Special care was given
to prevent condensation water from flooding the medium. The GrQbler's
ink was diluted twenty times with n/6 glucose solution and sterilised in
the autoclave. In a sterilized, empty petri dish ten drops of the ink were
put in a row. The first drop was inoculated with a small amount of bac-
teria from a twelve-hours-old streak culture. The bacteria were thor-
oughly mixed with the ink of the first drop. Then a loop of this was trans-
ferred to the second drop and also thoroughly mixed. This transfer was
repeated in the same way with the remaining drops in the dihh. From
the tenth drop, with a sterilized drawing pen, a small amount was taken.
Small (lots were made with the pen on a gelatin plate in such a way that
the surface pellicule of gelatin remained uninjured. These ink dots were
left to dry for two minutes then covered with a sterilized cover-glass. A
small drop of immersion oil was 8ul)sequently applied to the cover-glai«
and the whole pt^tri dish was i)rought to the microscope for examination.
Each black p<jint wa.^^ then examine<l \^'ith microscope until one was
found which containoil one single organism. The organism appeared as a
translucent dot on a black field. Its development was closely foUom-ed;
the first division was distinctly noticed after half an hoiu*; it muhiplied
rapidly; all the individuals were motile; they liquefied the gelatin riightly
and nioveil alK)ut ver>' briskly in the liquid under the cover-glass. After
six hours some of the organisms l)ecame sluggish and gradually k)st their
motion. They incn*a.'*<Ml in size and forme<l a capsule. Repeatedly they
divide<i in the same capsule, stretching the jelly-like membrane more and
more. The newly forme<i organisms within the original capsule also en-
ca|)sulated in their turn (figs. 0 and 7). At the same time the indi\idual8
on tlic rim of the colony multipliefi and remaine<i mcvtile. The double
form of the bacillus was in this way clearly explaincMl and pn>ve<l.
IDKNTITY OF THK OKCANIHM FOUND IN TURLY TOP BEETS
\ Study of the litcnitim' of the subje<-t shows that the greatest similarity
to our organism (»f any descrilN'd s|K'ci<»s is presented by that descrilieil by
.\nhur and Holl(*y^ as liacUrium Dianihi as the cause of a leaf spot of the
caniatic»n. In its morpholog}'. so far as descrilKxl by these writers, this
organ i;4iM is v(*r>' similar to ours, the n*semblanc<» lieing nmde pronounced
by thf (i«-veIopinent <if characteristic z<M)gl(H*a<'. In biological liehavior,
Imwrver. the iwo or^canisins cannot Im' accurately compared, since the
* \rttiiir. .) i' :ini| liiillfv. II. L. HiirtrriiMiiN (»f rnriintioriii. Indi»n» .\Kr. Kxp
St:i lilt I .v.* IVHi.
1917] Boncquet: Bacillus morulans 285
work of Arthur and Bolley was carried on at a time when bacteriological
technique was not standardized upon modem lines. One noticeable dif-
ference exists in respect to growth upon an acid medium, B. DiarUhi being
said to grow best under such circumstances, which is not the case with
our organism. The description of the bacterial organism given by Arthur
and Golden^ and again by Miss Cimningham^ as the cause of the so-called
Indiana sugar beet disease, is similarly subject to imcertainty, but if this
work was accurately done the organism must certainly have been differ-
ent from ours in that it is said to be a particularly active gas former, which
feature is totally lacking in our organism. It seems proper to mention
here, however, the fact that Professor Arthur in a recent personal letter
states that the accuracy of all this early work performed under his direc-
tion is open to some doubt on accoimt of the imdeveloped condition of
bacteriological technique at the time and he expresses the opinion that
the organisms found by Bolley, Miss Golden and Miss Cunningham were
very likely identical.
The organism described in the unpublished work of Schneider* as BaciU
liLS califomienaiSj which was isolated from curly top beets in California,
seems again in its morphological characteristics to be entirely similar to
ours and we feel little doubt that Schneider and the present writer had
the same organism before them. Schneider found his organism also very
abundant in sugar beet soils and upon the surface of the plants. He at-
tributes a stimulative effect to this species, when applied in pure culture
to sterilized beet seed or to the foliage of yoimg plants.
The organism described by Dtiggeli* as being abimdantly and often ex-
clusively present upon the surface of various plants and seeds, seems also
very similar to ours in form and size, formation of zoogloeae, color, sapro-
phytic habitat and most biological characters. This was named by Diig-
geli Bacterium herbicola aureum, but "said to be the same as the BaciUus
meserUericus aureus, isolated by Winkler from the surface of plum leaves."
The latter statement confuses the identification.
DESCRIPTION OF THE ORGANISM
Summing up the whole situation, we feel justified in describing oiu:
organism as a new species on account of the incomplete and doubtfully
* Arthur, J. C. and Golden, K. E. Disease of the sugar beet root. Indiana Agr.
Exp. Sta. Bui. 39, pt. 3: 54. 1892.
^ Cunningham, C. A. A bacterial disease of the sugar beet. Bot. Gas. 28: 177-
192. 1899.
' Schneider, A. The California beet blight. Spreckels Sugar Co. Exp. Sta.
Rept. 28: — . 1906. (Unpublished.)
• Centbl. Bakt. II, 12: 602 and 695; 18: 56 and 198. 1904.
286 Phytopathology [Vol. 7
accurate descriptions of those species which more or less rcsemblo it . ami
the fact that none of them corresponds throughout.
Bacillus morulans n. sp.
Morphology
Vegetative cells. Grovm in Liebig bouillon for sixteen hours at aUiut
20°C., oval to short rods, single or in pairs. Groim at 37*, shoft rrMb in
pairs or in short chains.
Sizi'. Length 1.5 m; breachh 0.1) n: extreme length from 1.5 to 2 m-
Capsules. Kasilv observed in 1 lOCK) glvcerin iKmillon after twenty-
four hours and also in milk nuKlia (figs. 6 and 7).
Motility. Very active on agar and in l>ouillon, when grown at 37' for
twelve hours.
FUujella. StaiiuMi by Z<»ttnow*s meth(Hl; four long peritrichial flagella
(figs. G and 7).
PU'omorphism. Cocciforius obstTved in glucose Inmillon tulles and
bloo<l s<»niin nie<iia after thirtv davs.
Stain. Miisilv with waterv fuchsin, d(H*olorize<l bv CSramV m€*thcNl.
Cultural features
(ielatin platv mutral to phenol phthalein, Konn. round to irregular;
surfa<*(» elevation. Hat to convex contoure<l; internal Mnieturv, refnuiion
strong, hyalin**, inoruloid; zoogl(K»ae very marke<l; tHlgi*s. entire to undu-
late: optical characteri.<tics transparent to butynnis; consistency. \'u*ctm.*.
Kacli colony is surrounded with many .^'condary colonies, appearing a.H
sniall. «»ily drops of high refra<'tiv<* power. The appearance of so(»gloeaf
is verv noticeable in the middle of the cohmii'S.
(it hit in plntfs I ..'t oeid tn phvnolphthalein. The entire nuM» is a xooghM^a,
IoIn'iI and irregular in outline; the colony is slightly colored; orange-yrl-
Inw. !in >iirroun«liiiu ro|nni«»s noti(*e<l.
fnhit'f, stnnk. After five days: gro\\lh, linear; margin, continuoiL«;
surf.-m- iriji-f, flat i<» eiuivex: li^Jht transmissi<»n. butynius; t*olor. yellow-
oraii^*-: lu-^trr. ^li-trninj:: cimsi^tency, visi-ous. The water of conik*n«&a-
tinii ha^ :i yrjinw >4Mliiiieiit .
ft't iitt. /i.'/». Aft IT twi'iity-twn h«»urs. top gn»wth:si£e. 5 mm., im^ilAr
('(iiiiciit*'d pulvinate tn capitatr. li^lit orangt* in color; vis(*ous in coiLM^t
<*in\ . 'ii-tti ^liiiiiim.
(,..>' / ^ti,ttl . I'ilifnrin tn slinlitly iM*adi'd. .\fter fifteen <lays. liquid
f.f Ml. ii «.i iiii'diuiii. rratrrifnrm with a vellow s4Miiment. After Iwenlv
iii>-. li«j»i« fretimi ytratiforiu; yellnw pigment. decreas<M| by alMtenci* of
288 Phytopathology [Vol. 7
Agar. The colonies are extremely variable according to the deniQty trf
growth, the moisture and the t-emperature.
Milk. Peptonisation of casein in fifteen days at 37®. The re«iclMm w
alkaUne to Azolitmin.
LitmuH xvhcy. Remains clear, alkaline reaction.
Bouillon Uibea. Opacity l)egins after eight hours at 37**, a pellicule
fonns in twenty-four hours or less. The color of the pellicule antl the
ring is dull, soft gray ; thick, viscous and conaists of oongkimerate leogloeiir.
The8e are generally oval, but may l)e linear and all united in ehainii.
Deposit, forms after two days incubation at 37**. I>epoKit is in the
lK*ginning slight, and finally yellow. The amount of deposit and the in-
tensity of the color increases, how<*ver, rapidly. After t*»n days, a de-
cidedly d(»ep yellow-orange has develofx'd. The de|MK<it is compact and
viscid on agitation.
Potato streak. After twenty four hours at 37**. Size, 2 nun.; sharp.
linear; margin, continuous; color, yellow, honuM*hn)mous ; lust<T, glisten-
ing; texture, homogencHnis. Xo Hcjuefaction of |)otat<» and no gat
formation.
Physical a tut biochemical features
Heaction. In carl M)hy drat <'-fre<» nuMiia the reaction is alkaline; in carlMH
hydrate media, the reaction is acid, except in lactos4», where the n*action
is .slightly alkaline. S«m» table <>.
\ Urate Liebig broth. After twenty-four hours at 37', stnmgly nNhice^l
to nitrite.
ludol. Not pHMiuaMi in p<*ptone solution after ten days.
Optimum temfnrature. 37**, meju^unnl by the amount of acid pnMiu<'«'d
in 1 JMT <'ent glu<*os4* after five days. Acidity w:l»* 2.5.
Thermal death fmint. Six-hours culture in iMmillon; i'A'^i \ in ten minui<*9».
Carlnth yd rate fermentation. Shown in table ti.
lirsistanee to mercuric chloritl. Six-hours <*ultun' on iKiuillon agar stn^ak
killc<l in 1 2:>.(NN) to 1 ;i(MMM) in ten minut<^.
Ut latter growth in acid and alkalin media. IX*tennin(H| by the ap|M*:ir-
aiKM* of cloudinos in the tulH>. (Sniws bc^t cm neutral or slightly idkalin
mrdia. Vwv \h't cent in ari<l appan'ntly stops all gnmih; 7 per ei-nt in
alk:tlin«': >am('.
<ffi> piiuhu'tnni. No gas i> pHNJuctMl. Sim* tabic G.
litlntittu tit frtt oiygtn. AiTobir; facultative anaerobic.
1917] Boncquet: Bacillus mordlanb
TABLES
CarbohydTate /ermentation of BaeiUui morulana
(Incubation: 37''C., medium neutral to aiolitmin)
NoTx. — A, acidi B, basic; p, permanent; b, abundantly present; a, absent; x,
more or less present; *, acid on top and basic in tuho.
PathogeneaU
One loop from a twelve-houre-old streak culture on bouillon ^ar intro-
duced intravenously in a rabbit, caused death within twenty-four hours.
On Diajithus incamaUi. The young unfolded leaves,' when unrolled and
covered with an abundant suspension of bacteiia, developed small necrotic
r^oDS. The necrotic regions are watery and translucent on the edges,
slightly elongate or irregular in outline, following the venation. The in-
side of the necrotic r^ons is sUghtly brown.
UmvBBsnr or Caufoknia
Bebkelet, Califoknta
A NEW APPARATUS FOR ASEPTIC ULTRAFILTRATION
Ralph E. Smith
With Two Fiquees in the Text
In various investigations of so-called nonparasitic or phyaological plant
diseases, as well as in numerous animal diseases in which the pmence
of an ultramicroscopio organism is suspected, the juices or body fluids of
affecte<l plants or animals have often be< n subjected to filtrmtion through
Berkefeld, (^haml)erlan(l and similar filters in order to remove bttcteria
and other organisms of microscopically visible size. While inocuUtions
with such filtere<i juice have lxH*n made frequently, and in some cases
(tobacco mosaic) with i)ositive results, the possibility of dcmonstniting
the presence of a t)nrasite in the filtered juice \iy its poosiUe growth
when all other organisms an' excludeil, se<*ms to have received little at-
tention so far as plant diseases are concerntHl.
Another object in pnnlucing an aseptically filtered juioe is that of
obtaining a st^Tile, unheat<Ml plant extract as a culture medium, as men*
tion(*<l by Mr. B<>nr({uet in another article in this number of Ph>'to-
path()log>'. In either cast' the juice may l)e varied in concentration or re-
action, any desinul substance which will pass through the filler may be
addtnl to it. an<l in many ways ilh4* may hv found in the investigation of
plant diM>as<* f(»r tisi^ptically filtennl and preserved or ooM-ffteriliaiHl
jui(M\s. Various devict^ for this purpose^ have l)een described but all
of them, HO far as the writ<T is aware, an* clums}' and of very doubtful
efficiency. Those in which the candle and the Uquid to be filtered
in 11 tulN* <iin*rtly aUive th(* receptacle or outlet for the filtered juice
esiMM'inlly uiidrsirahle. since the slightest leakagf* may result in contam-
in.HtiiiK the filtrate with the unfiltenHi juice and thus d<*feating the whcilc*
objtrt of filtration. HiiblNT stop|)ers an* alwa>'s op4*n to sus|Nci4m.
ArranK«inentH ju whieh the filtrate is caught in an opi*n or a cutton-
stopiMii ve>Ml and then iM>ure<i into the* cultun* tul)es in the open air
are eiTtaiiily far fmm safe. Those in which the liquid is forced thmugh
the raiiiilf fri»in the iiL-iide outward are obj«H*tionjible on account of the
quirk eujitii»K ovt r of the inner surfaer with sentiment , which, on the i»ut-
>ii\r. may Im- iimMly Imishid off. In the ap|)aratus illustrated hen* the
only )Hi>-.il>ility ni <*ontaiiiination is fri»ni the air and cannot occur fn>m
th< \ihfUti n tl juicf. Ill 4 it her words, it is abs<ihitely c<-rtain that nothing
292 Phytopatholoot (Vol. 7
in the line of tubing from the candle to the upper reservoir are wrapped
closely with cotton, extending beyond the ends of the rubber connections
and for about an inch down over the candle itself. Over these an* put
again still longer cotton wrappings and finally a layer is put on covering
the whole line completely from a point two inches bdow the upper end
of the candle up to the reservoir. The whole region around the fKUip-
cock is similarly wrapped with several layers of cotton, with a largi* plug
of the same material in the lower opening and a wrapping over this and
up arotmd the tube. It is the intention that all the joints shall be air-
tight and in this apparatus the number of joints and chances of con-
tamination arc less than in most similar devices. (Compare for example
the arrangement figured on page 65 of Marshall's Microbiology.) The
cotton wrappings are added, however, as an extra precaution.
After the apparatus has been put together and wrapped it b sterilised
in the autoclave. At the same time a supply of cotton-phigged tf-nt-
tul>o8 is Rterilized by dr>'' heat. The apparatus is then set up on a taMe
convenient to a gas burner and some of the juice to be filtered pourrd
into the l)eaker, taking cure not to allow the cotton at the upper end of
the candle to touch the li(|ui(l. The water vacuum pump is starteil.
using as little suction as po8sil)le, and allowed to operate until a supply
of the filtered juice has l)een collected in the reservw*. If the aurfare
of the candle iK'comes Uh) much clogged it may be cleaned with a soft
brush, but it is (l(*sirable In^fore commencing the filtration to remove as
much solid maU^rial from the juice as possible by filtering through cloth.
pa|MT and sand. As soon as enough juice has l)een filtered the cott4m
phiK dosing the lM)ttoin of the main tul)e is removed; one of the steril-
ized test-tubes is h<'l(l in imv hand, the plug n>moved and discarded, the
tiilx* thoroughly flaineil s<*v(*ral inches do^Ti from the top and then panst^l
up int4> the lowrr end of the ap]mratas ]N*h>w the outlet tube. With the
other hand the stoiw'ock is tum(*<l and some of the liquid run intoth«*
tulM'. Aftrr shutting off the cork, another tulH» is taken from the ))ask«-t
with that hand, whilt* the tulx* just RIUhI is withdrawn and its open enil
hiM in the f1:iine. Th<' ])luK of no. 2 is now transferre<l to no. 1. Nith
tiilMS ronstaiitly flaiiieii. no. 1 is laid aside and no. 2 filled, continuing
th»* pn»e«->s with as ipimy tnU-s as «lesired.
Tht apparatus fiKtirtMl is 14 inches long over all« and holds 200 rr. «if
li«|iii«l. DitTiTi-nt <izis may of r«iiirse Im» n»ade and it is ver>' pn»l»al»l«'
th.it •^nim «.f thu'H«' inten-Med in the matter may \h* able to suggt*st im-
pri*vi lilt iii^ ii)M»n tlie apjiaratii^i. A eontimious glass tulM> n^Khl U>
n- 'i iM'twirn thr riM-rvoir and filter eandle, thus eliminating tme juint.
l»iif It hii" In I II thuiiirht that the inerea*<eil liability of breakngi* n^uulti
m«>ri- than ntT<*et any disulvanta^ce. It n*ight also Im* {MMsible to rnnent
FACTORS AFFECTING THE PARASITISM OF USTILAGO ZHB
F.J. PlEMBIBEL
Because certain facts seemed to indicate a different life hitttorv fmm
that usually credited to Ustilago Zea (Beckm.) linger., on corn, field inocula-
tion eicperiments were carried on during the summers of 1913 and 1914.
The results of the experiments were such as to make a more detailed
investigation of the spores and sporidia highly desiraUe.
The works of Von Waldcheim (10), Brefeld (2, 3), Hitchcock and
Norton (6), and Clinton (4) have given us the salient poinis in the life
hudory of this parasite, and numerous other investigators have niaile
additional contributions from time to time, showing that the control
of the fungus is a difficult problem. The spores of the smut are wi<lely
distributed by the wind and are produced in large numbers throughout
the growing season. They are capable of germinating immediately and,
in a suitable medium, they produce immense numbers of sporidia which
may bud in a yeast-like manner and produce a host of others. The pro-
duction of the sporidia in largd numbers in the field is possible in such
places as manure or compost heaps.
Brefeld (2) ccmductini a few expc^riments from which he concludctl that
s|K)ridia arc short livo<l, d\ing in five weeks when dry. The sporidia
thenifore have l)cen chanicterizeil as Innng *Wiort lived" and ver>' little
is really known concerning the factors affecting their vitality. Nor do
wo know the fate of spon's on the com used for ensilage. The prrarnt
inv(^igation was made in an effort to secure more definite information
on thes4* jMiiiits.
INOri'LATIO.V EXPERIMENTS
Methods
Pun- (*ultun:9< of tho fuiif^us wort* ohtuimMl by the poured plate method,
u.*(u:illy on Ix'^Twort aK:ir. Tho c<)loni«'$i wen* later tranjiferrod to agar in
tulH*^. SiM»ri<iia from pure (Miltun* wrn* then used in the inoculaticin
ex(NTiiiii*iits. IiiociihitionH wt'n* ni:ui«' either by smearing thi* sporidia
din-rtly on in tin* plant parts or by |>la('inK th«>m in water and applying
\\i\> ^u^^M•Il•^if^n of >|H>riiiia by inrans of ji dn>p|H*r or hypodemiic s\Tingiv
'rh«* hypiMit-niiic >yrin^(' w:i> umsI when it was d4*sin*d to incKruIato the
vvrv voiinK part> which had not v«*t Ut^n unfolded.
1917] Piebieisel: Parasitism of Ustilago Zisim 295
Spores were ako used in inoculating the plants — ^usually they were dusted
directly into the tops of the plants or mixed with moist soil and then
applied. In a few cases the spores were applied in suspension in water.
Factors affecting infection
During the summers of 1913 and 1914 a total of 2064 plants of Minne-
sota no. 13 com were inoculated. In all eighty different series of plants
were used in the tests. The highest percentage of infection in any series
in the summer of 1913 was 70.8 and in the summer of 1914, 84.2. These
results were obtained by injecting suspensions of sporidia in water into
the growing point or as near it as possible.
Experiments were made to determine the effect of the following factors
on the success of infection: age of the plants; injury to plants; age of the
spores and sporidia. Observations were also made on the relation of
early planting close planting and soil conditions to the amount of smut
present in tha fields.
Age of the plants. It was found that successful infection depended very
largely on the age of the plants or plant parts inoculated. Healthy,
vigorous plants about 2 to 3 feet high are most susceptible. It was very
diflBcult to infect very yoimg or very old plants. These results confirm
those obtained by Brefeld (3) and Hitchcock and Norton (6).
Injury of the plants. Experiments performed by Chnton (4) showed
that mutilation of the com plants when about ready to tassel tends to
increase their susceptibiUty. The experiments of the writer Ukewise
showed that injury tends to increase the chances for infection. When
young leaves were injured and then inoculated, the resulting infection
usually spread from the point of injury as a center. Many of the infected
leaf areas, however, never developed sufficiently to produce mature spores.
This was probably due for the most part to the rapid maturing of the leaf
tissues and the consequent inability of the fimgus to spread through these
older tissues. Injury is not necessary, however, for successful infection.
Age of the spores and sporidia. The length of time that the spores can
withstand unfavorable conditions is important from the standpoint of
the propagation of any fimgus. Brefeld (2) demonstrated that corn smut
spores, eight years old, germinated, but he did not attempt to secure
infection with them. The writer obtained spores, five years old, and made
field inoculations with them by dusting into the tops of the plants. Six
per cent of the inoculated plants became smutted near the point of inocula-
tion. Two per cent of smut also appeared in the check plot but the
infected areas appeared on various parts of the plants.
Many fimgi when kept in continuous culture on nutrient media for
296 Phytopathology [Vol. 7
some time lone their power to infect. Brefeld (2) found that amut siporidia.
when kept for a year in continuous culture in liquid media, Iok( their
viability. The- writer kept the sporidia in continuous culture on lienr-
wort agar for eight months. The culture tubes containing the sporidia
were then buried in snow out of doors where they remained for a month.
Inoculations wore then made in the greenhouse on two com plants. (>no.
a snuill plant about 2 feet high, became infected in each of the five leaves
inoculated. The other, about 5 feet high, was inoculated on an ear and
prcxluced a large smut Imil. Thus it will Ik^ seen that continuous culture
of the sporidia on a solid nutrient medium for a period of eight month<«,
followed by freezing for a month, did not destroy their ability to infen.
Recently, portions of a pure cultiuv, that had Ihwu maintaine<l on l>eer-
wort agar sinct^ June, 1913, a period of three* and one-half years, were placiNl
in hanging drops of di.slill(Hi water and also of mo<Iifie<l ( ohn*s s4>lution.
In th<»se cultures individual sporidia wen* probably no longer present but
theR* was a miiss of short hyphal thremls, apparently resting segments
of th<* g<Tin-tulH^s of the sporidia. The ends of thes<* hyphal thn'iMls were
deiiw»ly pn)t()j)l:ihmic and wlM»n they wen* pUice<l in water, or, l>ett4*r
still, in a li(]uid nutrient me<liuiii they again pnNiu<*iHl sporidia by budding.
Tlu^M^ sporidia continucni to bud until the supply of nutrient matf^rial wan
ex)iaust(Ml when they again sent out long thin g<*rm-tul>es. The writer
inoculated four plants with some of the pure-i*ultun* material but has not
as yet succ<h'(Um1 in securing successful inftnrtion.
Kjftct of varly planting ^ char planting and soil conditions, Arthur and
Stuart (1) n*iM)rt that early planting, close planting and moist rich M>il
incn*ji^e the ani(»unt of .snnit. Il^iiny periods were also closely followeil
by outbn*aks of snuit in the com fields. The following obser\*ations
niadt* by the writer point to similar condusioiLs.
The corn us<sl for the inocuLition ex|H*riments was planted at succeMtive
intervals throughout the s<*ti.son .so as to have at all times an abundance
of i)lant.s in the sus(*eptiblc stage. The plots wen* carefully examined for
smut and it was found that a gn^ater amount of smut was present in the
plots planird carlitT in the M»2ison. The phuits in such fiehls are in tlu*
ni«»>t .suMTptible condition at a time when the weather Ls still fairly cool
and iMoiM ami the coiiditinns for infection ver\' favorable. The later
summer weather 'i> likelv to In* t<M»hot a n<i t<N)drv for successful infection.
I >:ite Mill in let with its I lion* frequent rains and c<N>ler weather again bring!«
on a fre.sh <iut)>reak of smut, the parts most aflf(*ct«sl lN*ing themdinientar>'
ears, siiin* tin? n*st «»f the plant is aln*aily t-o<» matun*.
W htii rtirii i> L!ri»\\ii in (*ln.s('ly plantiMi n»ws for gH*en fodder or silagp
purpo-i-. till- >inut i> aUu nmn* pn*valent. probably l>eeause the Mini in
such tirlils n'Miaiii** Mirruli'ut fur a longer iteritNi of time. In one such
1917] Piemeisel: Parasitism of Ustilago Zea 297
field which had been continuously cropped to com a number of years,
there was 25 per cent of smut as compared with 7 per cent in neighboring
fields where the com was 3 to 4 feet distant in the rows and on which
rotation had been practiced. Com on poorly drained soils or on those
which are too dry has less smut than that on a rich, moist soil. The
former types of soils produce weak plants that mature rather rapidly,
while a moist, rich soil produces a very vigorous succulent plant which
apparently is also more susceptible to smut. These observations further
confirm the conclusions of other writers.
Character of infection
Smut boils are often so generally distributed on a plant as to lead one
to suppose that the infection may have occurred on yoimg plants and that
the fungus then spread throughout the growing tissues. It is not im-
common to find a single plant on which the tassel, leaves and the primary
and rudimentary ears are smutted while adjacent plants are entirely
free of smut. It is therefore difficult to explain why the successive
infections should all have occurred on one plant. If the smut were sys-
temic such a phenomenon would be easily explained. Brefeld (3), how-
ever, concluded from his inoculations that the infection is local.
Field observations. In order to obtain more* complete information on
this point, the writer made careful observations on plants artificially
inoculated in the field. The results showed that, when successful, signs
of infection always appear in from ten to fourteen days after inoculation
and that the fimgus spreads but Uttle in the tissues. On several plants
where the inoculum trickled down the sides of the culm, smut boils were
produced at a number of points along the line, involving the leaves, the
primary ears and rudimentary ears. In nature, similar conditions might
easily arise when a spore falls into the water contained in the funnel formed
by the unfolding leaves and produces sporidia. The sporidia could then
be easily washed out to the various parts of the plant by rain. This is
especially probable since the spores of the smut can germinate as soon as
matiu-e and parts beneath the point of primary infection are particularly
Uable to attack.
Brefeld (3) and Kiihn (7) both obtained successful infection on a few
seedling plants but in all cases such plants were destroyed by the smut.
In order to further ascertain the results of early infection, ten very young
smutted plants in a fodder-corn field were selected and marked. All of
the plants were about a foot high and showed varying degrees of infection.
Of the ten plants under observation eight were killed by the smut in less
than a month and the other two were greatly stunted. These two plants.
298 Phytopathology (Vol, 7
however, produced healthy ears. Furthennore, many plants that were
found smutted when quite youn^ matured healthy ears.
Greenhouse inoculations. It han abeady been stated that in the field
the infection of verj' young com plants was ver>' difficult. Further inocula-
tions were made in the greenhouse upon germinating seeds. Forty-three
seiMllings were dipped into a water suspension of sporidia and then planted
in pots. None of the plants developed smut although they were allowed
to gnjw for one and one-half months. These result* indicate strongly that
infect i(m is purely local and not systemic.
THK VITALITY OF SPOKKS A.ND SI'(»mi)IA
It is a well-known fact that corn smut spon»s retain their viability for a
numlMT of y(*ars, U^ing ext namely resistant to unfavorable conditions.
V<»ry little, how(»ver, is known con<*(»nung the vitality of the sfMiridia and
their resi.^'tance to unfavorable con<litions has hitherto Ihhmi supi>oM'<l to
Yh" very slight. Th(i writ(T al.**o investigated the vitality of the siKin-s and
sporidia. The cfTcets of the* following factors on the vitality of >|H»n'^
wen' considertMl: (1) the silo, (2) t<»mix*ratun», (3) carlnm dioxiilr.
(4) acids, and (5) s4'asonal factors. Tlu' influence of the following fa<'-
tors on th<' vitality <»f sporidia w:is rcmsideriMl: (1) tem|MTatun*. \'2*
desicration, {'^) arids.
V italHij of spares
Since a considerable amount of corn .^mut must 1h» carritMJ over into
silos with the corn in the pnK*ess of .silo filling, it is imiMirtant to know the
fat" of such spon^s. T<J d(»t ermine this, si>on's wenM*olUM't<»<l inc|uantity
and phu'ed in several .silos for various h^igths of time. A brief sununar>*
of the n»sults is given Im»Iow.
Sufuplrs I. TIh^si' siK)n»s, which had just mature<i, wen* iHilhvt***!
S<^pt4'm)HT 11. HM4. eiu'los<Hl in a che<*s(M*loth bag and phuM^I al»out on«^
third of the way down in a wtMHien silo. A sample of th" .Nune lot w:lh kept
a> a rheek in the lalM)nitory at nM)iii t«»m|H»nitun*. Tin* sj)ore> utTi'
n^eovered from the .^^ilo on NovemlxT 2, 1914, after having Iki'Ii th«-r^»
alM»ut s« ven we<*ks. T\\r .^pon's w<in» fn)W»n and wen», then'fon». thawt^i
out gradually in the laboratory.
Satnph- II. TIm^' s|H)n*s w«»n' ('olUvtiMl and phu'e*! in a brit*k >d«»
S'pleiiilHT 2. 11M4. The s|M»n's wen* eiielosiMl in a bag and plaeeil alN»ut
l."» ii^i'.i I'rtim the lM»ttom of th<* >\\o and aUmt 2 fe«*t fn»in th«' M«le. Thfv
\\rr«- reioveHMJ M:ireli 4, nu.*>, aft«T having lKM»n in the mIo for tivor
t^ii'iilN-'ix wei»k.«». The s|H)n»s when nM'ovenMJ wen* frt>z<'n and then^fon*
the s:iiiipie wa> dividt^d into thret* {Mirtions :i> follows: (1) A |M»niou
1917]
Pibmeisel: Parasitism op Ustilago Zkm
299
was kept frozen; (2) a portion was kept moist at laboratory temperature;
(3) a portion was air dried and kept at laboratory temperature. The
object was to eliminate, if possible, the method of handling the spores
as a factor which might influence their germination.
Samples III and IV, The spores were collected, placed in bags and
buried in a silo October 6, 1915. Sample IV was placed about 2 feet
from the side and about 8 feet i^m the bottom of the silo. It was re-
covered January 20, 1916, and was kept frozen until February 8, 1916,
and then kept dry at room temperature. Sample III, which had been
placed in the center of the silo, about two-thirds of the way up, was re-
covered January 15, 1916. It was also kept frozen until February 8,
1916, when the spores were gradually thawed out and then tested for
germination.
TABLE 1
Results of germination tests of spores kept in silo
NUMBER
or TRIALS
SPORE LOT
TESTED
MEDIUM IN WHICH TESTS WERE MADS
PERCENTAQB Or
OEHMINATION
Sample I
11
Silo
Distilled water
0
10
Check
Distilled water
25
5
Silo
Modified Cohn's solution
0
5
Check
Modified Cohn's solution
90
6
Silo
Tap water
0
5
Check
Tap water
12.5
Sample II
42
Silo
Modified Cohn's solution
0
14
Check
Modified Cohn's solution
90 to 100
10
Silo
Distilled water
0
8
Check
Distilled water
5 to 15
5
Silo
Tap water
0
5
Check
Tap water
15
Samples III and IV
18
Silo
Modified Cohn's solution
0
Check
Modified Cohn's solution
95 to 100
10
Silo
Sterilized distilled water
0
5
Check '
Sterilized distilled water
75 to 95
7
Silo
Tap water
0
3
. Check
Tap water
50 to 75
Sample V
50
16
Silo
Check
Modified Cohn's solution
Modified Cohn's solution
0*
75 to 95
• One spore germinated.
300 Phytopatholoot IVol. 7
Sample V. This sample was collected and buried in a silo about 12
feet from the top and 2 feet in from the side on October 4,1915. It was
recovered March 16, 1916, thawed out in the laboratory and germination
tests were made immediately. The results of the germination tests are
summarized in table 1.
The table shows that with but one exception the smut spores did not
germinate after having been in the site. Samples I and II were both
tested at the time they were collected, when 68 per cent and 28 per cent,
respectively, germinated in water. The single spore whidi germinated
shows a spore may occasionally retain its viability after ha\ing been in
silage.
The death of the spores may be attributed to a number of possible
causes — (1) unfavorable temperatures, (2) gases produced during the
changes in the silo, (3) the acids and other chemical substances produce<l
by fennentation, (4) pressure. The effect of each of these factoni except
the last was tried.
The effect of iemiyerature. The exact changes which accompany the
fonnatioii of silage from green com arc but imperfectly known. Esten
and MjL»<on (5), however, have shown that silage fennents best l)etwp«en
75°F. and 80°?., and that the temperature never rises alx>ve 86®F. in
profMTly prtjpanxl silage, except in the topmost layers where destructive
fennentation occurs. Noidig (8) n*|K)rts a nuiximum temperature of 91'F.
Such toiniKTatures are not in thi»nist»lves sufficient to kill com smut spores,
since Stt»wart (1>) foiuid that a teinp(»rature of 52*'(\, for fifteen minutes
was n(*cessar>' to kill tho siK)rt\s when immonHHl in water. He also found
that expo.Hure to dr>' hoat In't w(»i»n lOo.SX'. an<l 1()G°(\, for fifteen minut4««
kille<l th<5 spores.
In a few tests ina<l(* by the writer some spon* withstood a dry heat of
103 (\ for fiv(» niinut(*s, hut tus a rule a temiwratuns of lOO^'C. for five
niinutcs is .Kuflici<*nt to <U^troy the gcnninating powers of the sfxires.
Fn't^zing tcnijM'nitun's do not w<*in to injure the s|M>rps as they Hur\'ive
our .sovurest wintrrs. Si>on*s, ston^l in a sImhI when* they were nien*Iy
shclttToii from t^now and rain but were exposeti to all the rigors of wint4T.
girnniiiatrd wrll in >pring. S|H)res that were frozen for short periotLn i»f
tinif and tlicii lasted for germination ap|)eanHl to have l)een Ktimulat4*d
by \\\v viM. Thr fxtremcs in t(Mn|)eratun^ which the spores cncount4»r
in tlir >iln, thrn:fnn% can have no intiuenr^' on their vitality.
Th» tjhtt nj tjnsrs. Littfc IS known of the gases produced in silage
formaiinii altlmuKh it i> not pmbablo that they would take an active part
in tli<* ilr.otnK'tinii of the >mut >|)ort*.*<. ('arlM»n dioxide is probably pn>-
<lii<'«-ii ill ^n*:iif>t ({uaiiiit y a.-* a re>ult of the fermentative action. A single
exiM-nm«'iit ux'^ made to <iet<*rmiiu! the eiTi*i>ts of the gas on tlie sponrs*.
I)r\' ^|M>res wen* pla('«'d in a bottlu and carlnm dioxide was passed in. The
1917] PiEBfEisEL: Pabasitism of Ustilaqo Zrs 301
spores were thus exposed to the gas for ten days, when germination tests
were made. The spores were not only imharmed but germinated much
more quickly than those of the same lot which were used as checks. At
the end of two days no difference was apparent in the germination of the
two lots of spores.
The effects of acids. It seems probable that the loss of viability of spores
may be due to the chemical substances produced in silage. Considerable
quantities of acids are produced rapidly , the maximum amount of each
usually being produced within two weeks after the silo is filled. The total
acidity of silage, according to Esten and Mason (5), is about 1.0 per cent
to 1.5 per cent, the principal acids in order of their importance being lactiCi
acetic and propionic. In experiments to ascertain the effect of certain
of these acids on the germination of smut spores it was found that a con-
centration of 1 per cent of either acetic or lactic acids or a combination
of the two was sufficient to inhibit smut spore germination. However,
spores germinated in a diluted sample of normal silage juice.
The above results are significant in that they indicate to some extent
what happens to the spores in the silo. The optimum for the germination
of spores is reached at some point in the rise of temperature which accom-
panies the formation of sUage. The germination of the spores is, however,
inhibited by the presence of the acids. Acetic acid penetrates rapidly
and kills plant tissues. Hence, it is not unlikely that the spores, which
are exposed to its action for a long time, are killed by it.
Seasonal factors. Fresh spores were collected from time to time and
observations made on their germination. The first tests were made in
the summer of 1913 and fresh spores germinated very readily in water at
room temperatiu^. The tests made in the smnmer of 1914 were more
complete, begmning with the very first smut spores produced in the field.
The first germination test of fresh smut spores was made Jime 24, the
last test on October 10. Fifteen distinct tests were made, the results
showing conclusively that fresh spores germinate readily in water. In
fact, fresh spores germinated much better than did spores from the same
lot kept imtil winter. The average percentage of germination for the
entire series was 42.8, the percentages in different hanging drops varying
from 0 to 85, a result often obtained when water is used as the medium
for germination. This shows that spore germination in water is some-
what capricious, thus probably explaining the conflicting r^ults obtained
by various investigators. The germination in sugar solutions and liquid
nutrient media, especially modified Cohn's solution is more uniform, 100
per cent of the spores almost always germinating. No difference could
be foimd in the germination in sterilized distilled water, distilled water,
tap water or rain water. Incubating the cells at 24^C. to 38"C. did not
seem to influence the rate or amount of germination of the smut spores.
302 Phytopathology [Vou 7
Vitality of sparidia
Methods. In the studies of sporidia, pure cultures of the smut were
used throughout. Spores were sown in poured plates of beenn'ort nfcmr.
In about two days the .spores germinated and the position of the colonies
was niarke<l. At the end of four or five days when the colonies were
about the size of a pinhead, they were transferred to beerwort agar slanU.
A nunil)er of other nutrient media were also tried, \iz.: carrot agar,
nitn)gen free agar, beef agar, oat agar and a synthetic agar. Of these,
beerwort agar was foimd to l>o the Ix^, although a good growth was also
obtaino<l on carrot agar.
Cultural characters and morphology. When the colonies first appear
th(»y are round, raistMj, convex, opaque, slightly shiny to dull, light ctvam
in color. -\s the colonies grow older, the e<lge Injcomes somewhat IoIknI
and irregular, and the surface l)ecomes convoluUnl, ridge<l, or sharply
papillaU*. TIm* color (i«^p(»ns with age, lKH*oming light lavender in uU\
cultures. The consi.st<»ncy is first soft and n)py, then Ixvomes mucibigin-
ous or rublxTv; or. when k<»pt inoi.Kt, but\ToiLs.
TIh» colonics consist of sjKiritiia whi<'h are abjointed fnim the si<les and
(K*casionally from the end of the promyc<»liuin. These s|>oridia are of
the same iiatun^ iis those produ(*cd l)eneath the surface in liquid nie<tia.
The sporidia pro<iu<*cMl in the air from a liquid culturo are small, sharply
fusoid and fairly thick wuIUmI. Th<\v are pnnluced in long cliains. Thosie
produceii within the li(iiiid or on solid nutrient mo<lia are larger tlian the
air cronidia, not as thick walU^l luid aro somewhat roumled at the endf.
They are pluin|KT, <*ontaiii mon' oil globuk*s and are not priNluceil in
such long cliains. The walls an^ a[>pan'ntly somewhat nmcilaginoua.
In continuous cultiux) the s|K>ridia pnKiuce hmg gorm-tubcs, the emb of
which an^ dens4»ly i>rotophismic and which, therefore, can become nvttng
si'gnicnts |H)ssessing all the pn)|K'rties of sporidia. The gemi-tulNy Imv
come much cntangl(*d and give the cultun^ its rublM*r>' consistency, wink*
the disintegration of the empty portions of the hyphae give it its muci-
laginous chara(*ter. \ smear from such a culturo dries almost instantly
and b<*<>onu^ brittle. While s|>oridia pro<luccil in culture may not In*
exju'tly like thos4* pnMiu(*ed in nature, still they must Iw ver>' similar to
thos4* which we iniiigine an* phnIucihI in such great abundance in manure.
Thrrmal rtlatitmfi. A cf)mi)lete understanding of the tliemial relatione
of com sinut >|M»ridi:i would not only \n* of value in throwing additional
light upon the phenomena (»f inf(*<'tion, but it would also show more clearly
the nptitiiiun conditions for the profmgation of the fungus. An attt^mpt
wa-* thenfon* in.ide to as<*ert.ain the minimum, maximum and optimum
t4-niiMr:itun*>. Both tlried and actively vegt;t4iting s|)oridia were tested.
At tem|N*ratun*s nmging from 2i) C (o 2rr(\ the aniall sporidia bud
1917]
Piemeisel: Parasitism of Ustilago Z^jr
303
profusely in' nutrient solutions. As the temperature is increased to above
26®C. the sporidia show a greater tendency to produce long, slender germ-
tubes. At 35°C. growth is somewhat inhibited and the cells begin to
show an increase in the number of vacuoles. Increasing vacuolation
continues with a rise in temperature to 40°C. where growth practically
ceases, while at 46*^0. the cells are no longer alive.
Attempts to determine the lowest temperature which the sporidia in
liquids or on solid nutrient media can endure gave negative results be-
cause they withstood the severest cold of the winter (about — 28®C.).
Alternate freezing and thawing, however, kills moist sporidia. Desic-
cated sporidia, on the other hand, were not only able to withstand severe
freezing but in some cases were not severely injured by alternate freezing
and thawing.
When exposed to alternate freezing and thawing, however, there seems
to be some injury, as no subsequent growth occurred in two out of three
tests. Smears direct from pure cultures were not killed by drying for
one day at 21°C. or for fourteen days at a temperature of from 7® to 9°C.
TABLE 2
The effect of .temperature on desiccated sporidia of Ustilago Zece, AU sporidia
dried at tl^'C.
TBBT
Km-
BBB
KUM-
BBBOF
TBIALB
NUBf-
BBBOF
DATB
DBIBD
TBMPBBATUBX TO
WHICH BXPOSED
TIMB
BXP08BD
MEDITJM FOB OEBMXMATION
BULTB*
1
1
1
-IC^C. to -7**
14 days
Sterilized distilled water
+
2
1
5
43**
15 min.
Sterilized distilled water
+
. 3
1
6
Alternate freez-
ing and thaw-
31 days
2 per cent sugar solution
^—
4
1
7
ing
-2*»C. to 3**
12hr8.
Sterilized distilled water
+
5
1
12
45'*
15 min.
Sterilized distilled water
+
6
1
16
28.6^0. to 3r
24hr8.
Sterilized distilled water
+
7
2
16
-7
12hrs.
Sterilized distilled water
+
8
1
18
54^C. to 55^
15 min.
Sterilized distilled water
—
9
1
19
40^C. to 50«
16 hrs.
Sterilized distilled water
^
10
1
20
-5^C. to r al-
ternate freez-
ing and thaw-
12 da3rs
Sterilized distilled water
+
11
1
20
ing
-5C^. to 1* al-
ternate freez-
ing and thaw-
12da3rs
Sterilized distilled water
—
12
1
20
ing
28. 5*^0. toSr
24 hrs.
Sterilized distilled water
-
• No growth — ; grew weakly ^ ; grew -f.
304
Phttopatholoot
[Vol. 7
But Bporidia dried six days and then exposed to alternate freering and
thawing for thirty-one days were killed.
Sporidial ameara direct on glaas ooverndipe from the pure culture were
unaffected by drying for sixteen days at 21"^. and then at 28 JS"" to Sl'X:.
for one day. Sporidia first placed in water, then dried for twenty dajrs,
also withstood the same temperature. Sporidia in smean were not killed
by exposure to frotn 40^ to 50^C. for sixteen hours, after drsring for nine^
teen days, but appeared to be killed at 54^ to 55^^ for fifteen minutes,
TABLE I
The effect of denceatxon on the tntality of eporidia of Uetilago Zem. AU te*U for ger»
mination made al il*C.
•
a
i
1
p
2
MKTBOD or FBBrAKATlOir
KUMBBa or DATS DBIBD
MBDtm vmM9 worn , f
I'
I
i
1
Grown in cells in HtO
3
Sterilised distilled 1
2 days. Slip dried
water
2
3
Spores germinating in
H,0 in cell. Slip
dried
3
Sterilised distilled ^
water
3
5
In 11,0 in cells. Slip
1
Sterilised distilled ! +
dried
water
4
1
Smears made on slips
12
2 per cent sugar tolu*
tion
+
4
1
Smears made on slips
50
Modified Cohn's so- ^
lution 1
4
1
Smears made on slips
G at 21*C. 31 at al-
ternate f reeling
and thawing
2 per cent sugar so- - —
lution
i
5
1
In watvr on slips
12
2 per cent sugar so- -r
lution
5
2
In water on slips
48
Modified Cohn*8 so- -
lution
5
I
In water on slips
20 at 21 T.
24 hrs. at 2H 5*-3rC.
Sterilised distilled 1 *
water
()
1
»Snenrs on slips
•*
2 per cent sugar so* +
lution
A
1
SiiirnrH <in slips
iA
Modified Cohn's so- -^^
lution
1
1
Siiifiirs on slipH
1 ir, lit 2r(\
Sterilised distilled -^^
1
'2\ hrs. at 2S 5* 31*(\
water j
' , 1
1
Siiictrs (»n Hlipf«
' 1 liny lit 2l*('.
Sterilised distilled ^
11 diiys at 15»-2()t:.
water j
s :»
In W!it4'r on nlipn
124
Mo<iifird CofiD's so- ' -
lution \
ilmiiicH vrry :tl>iin(iant grrniination; + abundant; ^ sparse: — no gsniii<
nation.
1917]
Piemeisel: Parasitism of Ustilago Zea
305
TABLE 4
Viability of sporidia of Ustilago ZecB when smeared on a cover slip and desiccated
in light and darkness. All germination tests made in sterilized distilled water at
a temperature of Hl^C,
»
NVMBKB or TRIAlJB
•
• CONDinOK XTNDSB
WHICH DBISD
DATS DRTSD
BBtnXAS
2
Tiight
1
+
2
Dark
1
+
light
u
+
Dark
IJ
+
Light
6
+
Dark
6
+
light
8
+
Dark
8
+
light
11
+
Dark
86
+
Light
87
+
Dark
149
+
after drying for eighteen days. The thermal death point of dried sporidia
is, therefore, probably about 54® or 55°C.
Moisture relaiions. In most cases the sporidia were taken directly
from the pure cultures on beerwort agar and smeared on sterilized cover-
slips. The sUps were then placed in steriUzed petri dishes and were
allowed to dry at room temperature for various lengths of time. In
other cases a water suspension was first made of the sporidia and drops of
this were then transferred to the slips and allowed to dry as above. When
the germination test was made a drop of the desired medium was added
and the slips were then moimted on Ward or Van Tieghem cells.
From the results shown in tables 3 and 4 it is apparent that sporidia
can withstand long periods of drying without serious injury. Sporidia
when taken directly from pure cultures withstood drying for 149 days at
room temperatiu^. Not all of these sporidia, however, remained viable.
Sporidia first placed in water and then dried seemed to be less resistant to
desiccation. Sporidia thus treated grew after drying for twenty days,
but not when dried for forty-eight days. The latter result may be some-
what misleading as the number of sporidia in a water drop is much smaller
than the number in a smear from pure culture.
There was no noticeable difference between sporidial smears dried in the
dark and in the light. Light, therefore, ib probably not very injurious to
sporidia.
The above results are not in accord with those of Brefeld (3) who found
that sporidia were killed when dried five weeks. Nor are they in accord
with the statements of Arthur and Stuart (1) who characterize the sporidia
306 Phytopathology [Vol. 7
88 ''short lived" and further add, "These are borne through the air m*hich
must be rather moist or the aporidia will be killed by drying/'
Chemical reUUions. Sporidia were placed in various solutions of arettc
and lactic acid, in com silage juice, and in a mixture of various acids in
such proportion as to approximate the composition of silage juice. If com
smut spores germinate when placed in a silo, it was thought that by mean^
of tests with various acids commonly produced in silage the fate of the
sporidia might be determined. The sporidia in each case were obtained
directly from pure cultures on beerwort agar and were transferred to the
solution from which hanging drops were then made.
Sporidia apparently c^n grow in 1 per cent acetic or lactic acid solutions
or in a mixture of the two. In the lactic acid the sporidia tend to pri>-
duce a greater numl>er of gcrm-tulxw which probably indicates that the
medium is slightly unfavonible for gn»i'th. The sporidia also grow mrll
in expn*sKO(l silage juice. It seems probable, therefore, that if spore?! do
gonninate in the silo, the sporidia may continue to live in the silo for some
time. \Vhothc»r long oxposiutj to the action of the acids would ln» detri-
mental or not was not detenninoil.
A more concentrated mixture of acids such as was u.^e<l in the silaite
acid U^t proYtnl to l>e deleterioiw to the growth of the sporidia. The
sporidia appeariMl starve<l and l)ecame greatly vacuolate^!, a comlition
which probably pnMMHJes their death. The results here obtaineii are
not in a<'cor<l with those obtained when silage juices itholf m'as usetl,
possibly owing to the lack of sugars or other nutrients in the silage acid
mixtun). The efTect of the traces of but>Tic ami propionic acids alone
upon the .sporidia hjus not yet Ikh^u determined.
SUMMARY
1. Th<» inf<*ctinn of com by ('stilago Zecr (Beckm.) Unger is purely
IcM'al; no cvidontM; of sv.^teinic info<*tion was obtained.
2. Whrn very young plants Ikn'oiuo inftK^twl they are often killed.
'^. Injury to x\\i* host plant. clos4» planting, ver>' early or ver>' late
planting, and gn»wtli on rich soil ixxv con<iucive to heavy smut attacks.
•I. ViKonnisly gn)wing plants, Uitwwn two and three feet high, ar^*
moM .»<n.MTptil)U' tc» .smut attack.
/). TImj >j>nn'> of !'. Znr can <*au.si» infe<*tion either when young or ohi.
SiH»n*.s ^t-nniiKitf' readily a.s simhi :ls matun^ and n*tain their viability fur
!*cvrr;il y«':ir>: infiMiion wa.s o)»taincd by inoculating com plants with
sjHin"* fivf v<*;ir> old.
«i. riif rorn->!nur fuiigiL> d<M»s not los4' its vinilcncv quickly mhen
grown Oh aii.iticial ni(»li:i.
7. Thi' .'-pon*'* of r. Znr. alnntst without exception, lost their viabiUty
after having In^i-n kopt in a silo f<ir a few weeks.
1917] Piemeisel: Parasitism of Ustilago Ziue 307
8. The factors causing spores to lose their viability in the silo have
not been determined definitely; it seems probable that the silage acids,
especially acetic, may be the destructive agents.
9. Sporidia were kept in pure culture continuously for three and a half
years, at the end of which they remained viable. Inoculation experiments
with the same material gave inconclusive results.
10. Sporidia were desiccated for about five months without seriously
impairing their vitality.
11. Freezing injures sporidia but little; alternate freezing and thawing,
however, is injurious to moist sporidia, less so to desiccated sporidia.
12. The optimum temperature for the budding of sporidia is between
20° and 26°C., the maximum at about 40°C. and the thermal death point
near 46°C.
13. Sporidia can germinate and bud in silage juice, but are injured in a
solution containing acids in the proportionate concentration in which
they occur in silage.
14. The ability of sporidia, as well as spores, to withstand unfavorable
conditions is very significant in explaining some of the facts in the parasit-
ism of U. Zece,
Agricultural Experiment Station
University of Minnesota
literature cited
(1) Arthur, J. C. and Stuart, Wm. Corn smut. Ind. Agr. Exp. Sta. Kept.
12: 84-135. 1900.
(2) Brefeld, Oscar. Hotanische Untersuchungen tiber Hefenpilze. Die Brand-
pilze I, Heft. 6: 67-75. 1883.
(3) Brefeld, Oscar. Untersuchungen aus dem Gesammtgebietc der Mykologie.
Die Brandpilze II. Heft XI: 52-92. 1895.
(4) Clinton, G. P. Smut of Indian corn and teosinte. Illinois Agr. Exp. Sta.
Bui. 57. 1900.
(5) EsTEN, W. M. AND Mason, Christie, J. Silage fermentation. Conn. (Storrs)
Agr. Exp. Sta. Bui. 70. 1912.
(6) Hitchcock, A. S. and Norton, J. B. S. Corn smut. Kans. Agr. Exp. Sta. Bui.
62. 189G.
(7) KtjHN, J. Uebcr die Entwickelungsformen des Getreidebrandes und die Art
des Eindringens der Keimfaden in die Nahrpflanze. Bot. Zeit. 32: 121-124.
1874.
(8) Neidig, Ray E. Chemical changes during silage germentation. Iowa Agr.
Exp. Sta. Research Bui. 16. 1914.
(9) Stewart, F. C. Effects of heat upon the germination of corn and smut. Proc.
Iowa Acad. Sci. 6: 2: 174-178. 1895.
(10) Waldheim, Fischer von. Ustilaginea. Jahrb. Wiss. Bot. 7, pt.1-2. 1869.
SCLEROTIUM BATATICOLA
The Cause of a Fruit-Rot or Peppers
William H. Martin
During the pasKt few years the writer's attention has been called a number
of times to a rot of ])epper8. Specimens were sent in from various sections
of New Jersey witli iniiuiries as to the nature of the trouble. Except for
a drying and shrivelling of the epidermis in the more advanced stages^, there
was little extenial evidence of the presence of the disease. (>n breaking
the fruit o|x>n, however, the interior was found to contain numen>u8,
snudl, l)lack sdcTotia which were also present on the seeil. Isolations
were made, and in all cases pure cultures of an organism resembling
Sclerotium Ixitaticola Taub. w:u* securt^I. The marked similaritv of the
two organisms hsl to exiKTiments to establisli their pathogenicity as m*ell
as to determine if the>' wen* identical.
The fungus was isolaltnl by :iseptically breaking the diseasetl fruit; biin
of tissue were i\un\ pi<*k<»d out with a steriliztnl nee<ile and transfernsl to
a pounnl plate of nutrii^nt agar. In most cas<»s pure cultures wen* mIk
taiiied at tlie first planting. Inoculations were nuule as follows: Healthy
p<*p{MT fruits were inimerscHl for ten minut(*s in a 1 to 1000 s<»lution <if
mercuric* chl(»rid, w:i>hed in sterilized, distilkni water and place<l in a st<*r-
ilized moist rhamlKT. .\ttempts to pnxluce the disease by placing bits
of the rultun* iiiodia together with sderotia <m the unbn>ken surface
failed. Whitr tufts of mycelium were fornuHi but the fungus Mvmed
unabh* to |M'nrtrat(* the ei)idermis. However, inoculations on a cut >ur-
faci- iii:id(* with a fiaiiuMl sralfN^ were uniformly successful. A larg<* num-
Ut of iikoculatioiiv were nuule in this manner and 100 p(*r cent inftN-tiim
rr^ultrd. liifrrtioii was rvideiit in fmm four to si»ven days. Tho rpi-
d«Tini> at thr point of inflection Urame blackemni and the n>t >pn*aii
throimhoiit the intiTinr. AftiT thr fungus I NH*ame established, its pnign-sv
wa> rat I If T r:ij>id; in twn \V!t'k> the t»ntire interior of the fruits wen* in-
vailnl ah<l nunirnniN Mlrn»tia d«'velo|H»4l in the tissue and on tin* m^**!.
K\rr]»T lot :i M:irkt*ning of tin* I'pidiTmis there wen* no external signs to
in<li«-:itt' iltf |>ir^iii(-«> of thr di><'a>r. Inoculations wen* also nuuk* <in
Iniii -nil aTt:ii h««i tn thr pliint and in evrr>* c:iS4» the n*sultii were |Mte^itive.
.\ • iiip:> ut-rr ni.'nir t<> innnilatr thr roots ant I stems of growing plant>.
The Mill \\:i> rari'fullv rrni<iv<>tl from around the roots and an incbtion was
1917]
Martin: Fruit-Rot op Peppers
309
made with a sterilize^ scalpel and sclerotia from young cnlturee were in-
serted. The soil was then replaced. Roots similarly treated, but not
inoculated, were used as checks. Stems were likewise inoculated and the
cut surfaces were wrapped with cotton. On the stems and roots the inocu-
lations were not as successful as on the fruits. Where infection did occur
the fimgus was found just beneath the epidermis or in the pith. Death
of the plant or plant part resulted before the mycelium had advanced far
from the point of inoculation.
Reisolations were made from both the fruit and stem and the fimgus
was again grown in pure culture. Inoculations with these reisolated cul-
tures were, in every case, successful.
During the process of the above inoculations, similar inoculations were
made on peppers with cultures of S. bataticola Taub. isolated from the
sweet potato. In every instance they were successful. Not only was
this true but the characteristics and growth of the organism were, in all
respects, like the pepper Sclerotium.
In order to more fully prove the identity of these two pathogenes, the
following comparisons were made: (1) Growth on culture media, (2) cross-
inoculations on sweet potato and pepper as well as other hosts, (3) develop-
ment, measurements and the external and internal appearance of scleS^rotia.
Growth comparisons were made on com meal agar, potato agar, bean
plugs and nutrient agar. Abimdant growth was made and no differences
were observed.
For the cross-inoculations three strains of the organism were used.
One was isolated from the sweet potato, another, secured from Dr. J. J.
Taubenhaus, was likewise isolated from the sweet potato. The third was
isolated from peppers. These cultures may be designated 1, 2, 3, respec-
tively. The methods employed in the inoculations were the same as have
been previously described. The following inoculations were made:
TABLE 1
ReaiUts of inoculations on varioita hosts with three strains of Sclerotium haiaticola
HOST
Sweet potato (Ipomoea batatas)
Pepper (Capsicum annuum)
Tomato (Lycopersicum esculentum)
Cucumber (Cucumis sativus)
Apple (Pyrus malus)
Eggplant (Solanum melongena)
Turnip (Brassica campestris)
Red beet (Beta vulgaris)
Parsnip (Pastinaca sativa)
Carrot (Daucus carota)
NUMBBB
INOCULATBD
NUMBBB or
PXB CBNT
WITH BACH
CHECKS
INTBCTION
CULTUBB
25
10
100
25
10
100
3
2
100
3
2
100
3
2
100
1
1
100
3
2
100
3
2
0
3
2
0
3
2
0
310 Phytopathology (Vol. 7
In those cases where the inoculations were successful, no differennv in
gro^^iih of the organism were evident. The time before infection is e\-i-
dent, varies, however, with different host plants. On pepper, cucuiuImt
and tomato from four to seven da>'v is required, while on sweet potato,
apple, turnip and eggplant, from three to seven weeks elapses.
The development of sclerotia was followed in Van Tieghem cells. Tlie
method employed wa^ to pour enough of a melt<ed nutrient medium on a
cover glass to form a thin film. Aft-or this had cooled, it was inoculateil
and inverted over the glass ring. If care is taken in this procedure no
contamination results. By this method it is easily poflsible to trace the
development of a single sclerotium under high magnification of the
microscope.
Some cases were not^nl where sclerotia appeared to have developer! from
a single mycelial strand; in the majority of cases observe*!, however, sev-
eral or a numlHT of strands were involved. Sclerotial development is
pre(»ed(Ml by the formation of many s<*pta. Short mycelial tulies are
formed which ('onniM't the strands. These in turn become ver>* much
septate, as do the radial hyphae which are forme<l laUT (fig. 1). As tlie
wlerotiuin enlarges the central cells IxH'ome ccmtorted and fonn a ctMn-
pact' mass, due p(Thaps to the pn^ssitfe or resistan<*e of the out4*r cells
(figs. 2, 3, 4). The mature sclerotia are coal-black in color and in nuirt
cji.**<»s are fnM» fn)m any surface irregularities (fig. 5). No differences wrre
olx^Tved in the formation or color of s(*lerotia of the different strains.
In tlM» progress of X\u*si* studies numerous instancies were oIw^tvih! wIhtp
my<*elial branches united: this took place U'twevn bmnchert of a single
strand tis well as lM*twcM»n two s<*parate strands (fifp*. G, 7, 8). In stmie few
ca.*4<»s, this was the first i4ep in sclerotial development, but usually tlie
mycelium U'comrs M'ptate and irregular.
Sections of the srlcrotia of the different strains showed them to U^aUke
with H'gard to tlirir internal struct unr (figs. 9 and 10). In this conntv-
tioii s4HMis wen* S4H'iione<l to determine if the mycelium pi*iu*trat4\i the
m^mI inat, liut in nonr of the viim^ exjimine<l was this ol»siTve<i to U» tru«'.
The apiM'aranre of x-lerotia on the s4»<mI did not ap|)ear to afT«H't germina-
tion. Ilralthy plants wi»n» grown from disc»;iscHi setnl. Measurement.-* tif
xUr M-lt-nitia gave nvMihs identical with thos4» of Taubenliaus.'
Atti'iiipt> to produti* a jHTfecl .»<tage faile<l. In ever>' case, on the ger-
mination i»f tin* x-lrrotia ih'W sclrrotia wen» formed.
* J J T:tiiNnh:iUh The Murk rntH iif th«* ffWiH't potato. PbytopAth. S: iy> \%\
19171
Martin: Fbutt-Rot of Psppers
SCLEROTIAL DeVBLOPUENT
FiOB. 1, 2, 3, 4 and 5. Stages in Brterotial development.
Figs. 6, 7 and 8. Anastoroising of hyphae.
Fig. 9. SectioDS through mature sclerotia.
Fig. 10. Section of Bclcrotium on epidermis of pepper fruit.
312 Phytopatholoot [Vol. 7
conclusions
The pathogenicity of Sderotium sp. causing a rot of peppers {Capsicum
annum L.) has been established.
The following facts warrant the conclusion that the sderotiuni occurring
on peppers is identical with Sderotium baiaticola Taub.
a. CroHg-inoculations on sweet potato and pepper as well as other horts
gave positive results.
6. With both strains the growth was identical both in culture and on the
host.
c. Measurements of the sclerotia are identical.
d. Sclerotial development, color and structure is the same with liotb
strains.
It appcjirs fn)m these studies that the charcoal rot of sweet p«»tat4i«'!*
(Jinmiaa batata) is common and widely distributed throughout «H'tion«i of
New Jers<»y, that S. Imtaiicola Taub. can probably |x»rj*L»<t on several nthor
hosts and that it is the (*ause of a di^e:u<e of minor importance of p<*p|M*i>.
Laboh.\T()KY ok Plant Patiioi/kjy •
New Jkicsky AGKurLTiRAL Expeiument Station-
New PurxswicK, New Jerskv
A NECTRIA PARASITIC ON NORWAY MAPLE
Mel. T. Cook
During the suimner of 1913 the writer's attention was called to an in-
teresting disease on a group of Noirway maples growing on the private
grounds of one of the residents of Princeton, N. J. The first symptoms
of the disease were the wilting of the leaves and dying of branches very
similar to the well known symptoms of the chestnut bark blight disease,
caused by Endothia parasitica. In fact, it was the very striking resem-
blance to the chestnut blight that attracted the attention of the superin-
tendent of the grounds and led to the writer's being called to make an
examination. A further examination of the dead branches showed a still
more marked resemblance to the chestnut bark bUght; the dead branches
had been completely girdled by a canker which showed a blackening and
slight sinking of the diseased bark. The older cankers were covered with
the orange-colored pustules of the Tubercularia stage of a Nectria and
there was abimdant evidence that this organism was the cause of the
trouble.
' The superintendent had been watching the disease for some time and
had removed many dead branches and cankers. Fresh woimds were very
quickly attacked by the fungus, which made its first growth in the oozing
sap, but the careful treatment of these wounds with antiseptics, followed
by an application of paint, greatly reduced the number of infections.
The breaking of the small lateral twigs from the trunk and larger branches
was the most common source of natural infections and most of the cankers
had started from woimds of this kind.
An inspection of the trees on the estate and in the immediate vicinity
showed two others badly affected. These were destroyed, but the clump
previously referred to was left standing for observation and study. Dur-
ing the remainder of 1913 the infected parts were removed as soon as de-
tected and the wounds treated. In the fall of 1913 and the spring of
1914 heavy appUcations of fertilizers were apphed to all the trees and ob-
servations continued during the smnmer of 1914. The disease reappeared
and the fungus was always presentr on the cankers, but was much less
severe than in 1913. The fimgus was frequently found in the dead bark
around old wounds, but in many cases did not appear to be parasitic.
About one-third of the trees in this clump showed the fungus in 1914.
Another clump of trees on the opposite side of the driveway showed but
314 Phytopathology (Vol. 7
vcr>' little of the funf^us. The clump of trees has l)een kept under <»lKM»r-
vation (hiring 1015 and 1916, and the disease, although present L*« much
less severe than in 1913. Wounds are frequently infected but the in-
creased vigor of the trees, and the careful removal of diseased hranche?*
as soon as detected, has apparently resulted in a great re<iuction (if th«'
disease.
Since 1913 the writer has frequently found the fungus on Norway mnpl<^
and also on the muUxTry, working saprophj-tically and also appariMitly
as a weak parasite.
The idea that Xectria Is parasitic is not new in either Kumpe or America.
Weluner* reports having found it on the healthy stumi)s of a cut -over thi(*ket
of tnH»s and shruh.*^, esiKiciiilly on Cnrpinus sp. He also found a I>ipl«Mh:i
and a TulMTcularia on th<» young twigs of walnut* som«»times associ:it«Hl
an<l sonu»tinies the Tul>ercularia growing alone. He states that the twig>
wen^ winter-kill(Mi, and that the fungus gained access to the host thn>ugh
the dca<l parts from which it worke<l its way into the living tbisues. Tho
injunnl twigs w(T(» d(»foliat<»d and the tnn* lK)re vcr>' little fniit.
It was al.*«o reported !)y Hehrcn.s- as attacking Abies iHilsamnt, llie
terminal buds wcn» fr(^<|uently swollen. This wjis due to thr fomiati<»n
of a laver of <*ork lH'tw(»en th(» healthv and necnitic tissues. Tlir >\iell-
ings of the ])r(^vioiis year wen* mo.»<tly dead. Some of the gn^Mi twig> diii
not develop their l>ii<ls and .•<ome died later. Th<' myct*lium hilN'rnat4*<i
in the dead wood and |)<Mietrat<'d the living wcmhI the following .*«4*:lm)ii.
Thr mo>t important .Vmerican rejxirt is "by Polh»ck' who found Xtctna
CiKcinm (Pers.) Fr. causing cankers on yellow birch. Thes<» cank<T> fre-
(jucntly ginllc the infected parts. In cas<» <>f inftvUMi twigs hy|M*rtn>phics
w<'n* frMiueiitly fornuid. Sevctral Am<Tican students have rf*|>ort4Hl >tmi-
lar observations to the author.
Atiitn ri/n KVL Kxpkuimknt Station
Nhw HHrN>wirK. Nkw Jkii.skv
» Wf-hintr. (' Zuni PrirrfitiMiiniH v<i!i Nrrtriii rinnjilmrinn Fr. Z«*it. Pflniitrnkr.
4:71 s| 1V»I
' |i<l»r«'n- J. Kill iH-iiHTki'iiKWiTtrH Vorkniiinicti von "NiTtria ritiiiabannA"
iiritl till' \•Tl»^l'ltllIlK^•M^lHl• ilif*it'M Pil/c}«. Zt'it. PtlHiizriikr. 6: 1*.K{ \\Ps. Ivj.'i
■riiliiMk. J. IV A <-:ttiktr (III vi'llow hirrh ami :i NciMriii aumM-inlod with it
I{«-|)t Mull. Ar.'ui Srifiiif 7: 1*.K).V
PHYTOPATHOLOGICAL NOTES
Notes on Razoumofskya campylopoda. Berries of the false mistletoe,
Razoumofskya campylopoda (Engelm.) Piper, growing on Pinus sabinianaf
were collected by Dr. E. P. Meinecke in the San Rafael Mountains, Santa
Barbara County, California (Forest Pathology No. 17026), November 12,
1914, and sent to the writers. Seeds from these berries were used No-
vember 19, 1914, for inoculating yoimg pine trees in pots as follows:
one Pinus hanksiana Lamb., one P. bungeana L., four P. caribaea
Morelet, four P. contorta Loud., two P. couUeri Lamb., one P. densiflora
Lieb. & Zucc, two P. halapensis Mill., six P. mayriana Sudw., two P.
monophylla Torr. & Frem., one P. nigra Arnold, two P. parvifiora Lieb. &
Zucc, six P. pinaster Ait., one P. pinea L., one P. resinosa Ait., two P.
rigida Mill., four P. sabiniana Dougl., one P. strobu^ L., and two P.
virginiana Mill. Two trees of Larix ocddentalis Nutt. and two of Pseu-
dotsitga taxifolia (Lam.) Britton were also inoculated. The trees used
were from three to six years old. The seeds, enclosed in pxilp, were placed
chiefly in the axils of the leaves on the younger portions of the shoots,
adhering firmly as soon as the, pulp dried. Many germinated, but the
radicles of only a few succeeded in penetrating the bark of the trees on
which they were borne. In six months plants became established on the
following species of trees: one Pinus banksiana, one P. bungeana, one
P. caribaea, one P. pinea, two P. sabiniana, and two P. virginiana. On
Pinus bungeana and P. virginiana, dense witches-brooms formed around
the mistletoe infested region. On the other species spindle-shaped
swellings without witches-brooms were usually produced at the point of
attack. All these trees except Pinus sabiniana are new hosts for this
species of mistletoe in this country.
All the trees inoculated successfully produced clusters of mistletoe
plants in 1916, none of which produced mature fruits, apparently owing to
lack of fertilization. In 1917 mistletoe plants are again developing on all
trees except those with dense witches-brooms.
The effect of the mistletoe is to stunt appreciably the growth of all the
trees inoculated, as compared to other similar trees of the same species
not inoculated. Only one of the trees successfully inoculated has died
from this effect after two years' growth, one of Pinus virginiana with a
witches-broom. In case of trees of the same species inoculated imder
similar conditions with Peridermium cerebrum Peck, and P. harknessii
316 Phytopathology (Vol. 7
Mooro, nearly 50 per cent of the infected tnH»s diinl inside of two yrars,
indicating that the stem rusts are much more clestructive t4> young pim*
than the falw* mistletoi»8.
Th(» mistletoe is a western species which grows \igorousIy on ea»*t«m
spiK'ies of pines. Since the infect e<l areas on young pines may not !>*•
conspicinais during the first w^ason's growth and l)ecause of the fact that
the aerial parts of the mistletm* ])lants are annual, and an; not usually
observed on donnant trees, insjM»cti(m of nurser>' st^ick is nt)t sufficient to
insure its freedom fr(»m this harmful parasite. Shipments fn»m the
Roeky M<»untain and Pacific regions to thos<' farther east should hi* fli-*-
couraged* as they are likely to carr>' the mistletoe, even though they may
ap])ear clejin. (hir east<*rn pines are at j)res<*nt fn»e from mistlet<N\i. and
should remain so.
(Iko. (i. IlKIMit'CMK AND N. HkX HiXT
Tht prtufuctinn itf spons by AUiTfiaria Sobini in purr culiurr, Thi?*
fimgus has Immii the sul»JM*t of s])e<-ial stu<iy l»y thf auth(»r at th«' I'ni-
versity of Wisc<»nsin during the i)ast three yrars. The seiireity of ••pun*
pnxhiction in jHin* rultures, which has \HH*n not**^! l»y pn-vitnis work#r«.
was at th«* nutst't a hindnnu'e to iuo<'ulation experiiiM-nts. TriaN with
twrnty difTrrt'iit kiii<ls of iiuMlia including some tr>ts <in thi* fffti't nf
variation «»f :u'i<lity :iud tt'inpfrature wen* iiiadi-, hut thr n*Milt w.i«*
always tin* samr, frw Np<.n's luing pmduefd. In Ffl»niar>'. llU.'i. ••\-
pt-rinirnts wt-rr Ingun in whi<"h the moistun* enntmt of thi* mrtlium anil
th<- huiridity «»f thi' :ur al»«>Vf 1h<- rujture wrre varit-d. Thi** lik«-\%i'^-
was without ri>ult> rxrrpt whui tin- eulttin* was shrt'ddrd antl thr iii> rt-
hum srvmly woiunltd. Tinlrr >urh tn*atmi*nt ennrmi»us *i|Minilatioii
w:i«» alway> mtup d ifig. I. liK A f«'W hjMjrrs wtTi* foriiird whm th»-
invrt'liiini aloiu- \va> eut. Thr iiio>t sur<'i'ssful lUfthod eonsi>ts in gn»w-
iiig tJif Alti-rnaria m pttri dish eultun*, on liard ix»tat«» agar, for ti n t«»
t^\» Ivi I lav-, till II, lir^t, »»hn'dding thi- ag.ar to liits (fig. 1, ^4 » and >tirnne
ti» -I parati- aii«i i v«nly diMrihutf tin- pitrrs; s<*<'ond. contniHini: f«'r
t\v«nt\-fnur 1m fnrtv-iiuht honrs tlnnaftrr thi* mojMurr nlation ^. that
jiarti.il ilr>inu out i< itY«rt«'«i without :dli»wing tht* more exposed -urfai • -*
Tc Im ( i.rcr i:(*-i(iatt «i and hai»l. 'V\m- lattrr ol»ji-<*t is acrompli*ih(*«l imi-t
n a«:i!\ l\ r« ri.iviiii: t l:i lid and i \|M'-iiii; thi- di<h t4» sunlight in a >tinliz« d
riji !-• < 1. .1/1.1 r I'T 1h]1 i:ir. It » vapi»rati«»n i- to«i rapiil. on-a'^ioiud :it<-
II i/mi: ui»!: -ttiili/ii! \\:ttir i- n« ci «»-ary. Spon-s W(*re «il'taintil \*y thi*
J- • t : I d III t.»:tl iliiikii' -- in »!n iriiul'atur at *J»i ( '. hut tIomt att«iitii*n
\\ : r« 'iijii' ■!.
*'*.-!?:• . uT jii.l t \]i' -I •! -nrf:ii t - ni thi :iuar thi ndivrlop- fn»m th« "-id
ri.'''!iir' t ii'U\..ik « t" rl..-i|v -t ptati-. t l-.iik-wallrd hyphar. from whii'h
1917] Phytopatholoqical Notes 319
Phytaphthora infestans, causing damping-off of tomatoes. Phytophthora
infestans has been recorded as occurring on tomatoes by both American
and European pathologists. So far as literature on the subject is avail-
able to the writer, no one appears to have noted this organism as causing
damping-off of yoxmg tomato plants. The following brief accoimt of a
severe outbreak of damping-off of tomatoes caused by Phytophthora
infestans may be of interest therefore to pathologists.
In June, 1916, a number of young diseased tomato plants were received
by the Ontario Agricultural College from J. W. Noble, Essex, Ontario.
In the letter accompanying the plants Mr. Noble stated that thousands
of late tomato plants in that district had been destroyed. The affected
plants were first observed shortly after the tomatoes were set in the field.
A glance at the plants revealed brown lesions and constrictions on the
stems near the surface of the groxmd. Many of the plants showing these
symptoms had fallen over, due to the collapse of the stems at this point.
On examining the stems with a hand lens a white fimgous growth was
clearly seen on the lesions. This when examined \mder the microscope
proved to be the conidiophores and conidia of Phytophthora infestans
(Mont.) deBary. After this all the plants were examined very carefully,
and on some of them the same fimgus was found, apparently causing a
blighting of the leaves; but the chief damage done jbo all of them was by
the destruction of the stem near the surface of the ground. The falling
over of the plants by hundreds in the field was what first brought the
disease to the attention of the growers.
Some idea of the severity of this outbreak of damping-off of tomato
plants may be had by considering the fact that out of 288,175 tomato
plants supplied by the Heinz Pickle Company, Leamington, Ontario ,
to Pelee Island growers, only 45,000 reached maturity. At least 50 per
cent of the plants that did not survive succumbed to damping-off due to
Phytophthora infestans.
It is interesting to note that weather conditions during June were
exceptionaUy favorable to the spread and development of Phytophthora
infestans, the rainfall being much above the average for June, and the
temperature relatively low.
J. E. HowiTT
State and National quarantines against the white pine blister rust. The
following table shows the State and National quarantine action taken to
date against the white pine blister rust. The action is so varied in char-
acter that it seems necessary to present it in this form. Similar action
is under consideration in a number of other states.
320
Ph ytopatholoo y
(Vol. 7
arses.
UTAT*
WaiTB riMBS
OSOMrLAS-
lA
DATS
QCASAMTIXBD ABBA
CJanadfi
All
None
November 14,
All foreign countries
^^^^••••M^4M ••••■••••
1914
United States..
P. strohus,
None
September 16,
Great Britain. France,
monti-
1912
Helipum. Holland.
cola,
Denmark, Norway.
laml>er-
Sweden. Kuasia. i^wrr*
tiana,
many, Austria, Italy.
cembra
Switserland
All
None
May 21. 1913
Kurope and Asia
All
All
March 10, 1916
Canada and Newfound-
land
All
All
June 1. 1917
All points east of. and
including the states of
Minnesota. Iowa.
Missouri. Arkansas.
I^uisiana
AH
U. ni-
fcruni
June 1, 1917
States of New Kngland
and New York
None
All
Juno 1. 1917
Kurope and Asia
(California
All
All
February, 1917
F^ist of Mississippi Ki%'er
Delawan*
All
All
March 2. 1917
All |K)ints outside state
Idaho
AIL
All
March 1, 1916
New IIam|Nihire, Ver-
mont, Massachusetts,
•
Connecticut, New
York. Pennsylvania
Indiana
All
All
March 13. 1917
All |M>ints outside state
KanHaa
All
All
March 10. 1917
All |Miints outside state
MaMuichuBctta .
All
None
June 1. 1912
Kun)|ie
MicluKAn
All
All
March 19. 1917
All |M>ints outside state
Minnesota
All
None
Apnl 30, 1917
Maine. New Hampshire.
Vermont. Maasarhu-
si*tts. RhrMie Island.
Cofinertimt. Nrw
N ork. N«'W JvTm \ .
Pennsylvania, uhi<i.
Wisconsin
Montana
All
All
July 17, 1916
New llampshirr. Vrr-
mont. Massachusetts.
(Connecticut, New
York. Pennsylvania
Nrvftd.n
All pines
1
1
All
1
March 10, 1917
Fast of Mississippi
Kiver and Minnesota:
all foreign countries
Nr*» Jrrwy
All
1
1
1 NolM'
April 16. 1917
Maine. New llamfishire.
Vermont. Massachu-
setts, Rhode Island.
Connect tcut. IVnnsyl-
vania. New York.
Minnesota. Wisconiun
1917]
Phytopathological Notes
321
BTATB
New York.
Ohio.
WHITE PINES
All
RIBE8,
OROSSniiAR-
lA
None
DATE
March 24, 1917
Oregon.
Pennsylvania. . .
South Dakota^...
West Virginia...
Wisconsin...'
P. strobus,
None
monti-
cola,lam-
bertiana,
cembra,
excelsa,
flexilis
All
All
All
None
All
All
All
All
P. strobus,
None
monti-
cola, 1am-
bertiana,
cembra.
excelsa
February 21, 1917
QUARANTINED AREA
July 24, 1916
March 12, 1917
April 3, 1917
April 18, 1917
June 1, 1916
Ohio, Indiana, Min-
nesota, Wisconsin,
Maine, New Hamp-
shire, Vermont, Mas-
sachusetts, Rhode Is-
land, Connecticut,
Pennsylvania, Illi-
nois, New Jersey
All points outside state
East of Mississippi
River; all foreign
countries
All points outside state
All points outside state
All points outside state
All points outside state
Perley Spaulding and Roy G. Pierce
Personals, Arthur S. Rhodes, assistant in forest botany at the New
York State College of Forestry, Syracuse, New York, has been appointed
assistant in the Office of Forest Pathology, Bureau of Plant Industry.
Miss Ruby J. Tiller, scientific assistant in the Office of Forest Pathology,
Bureau of Plant Industry, has resigned her position to become the wife
of Prof. S. F. Acree of the University of Wisconsin.
Prof. L. H. Pennington of the New York State College of Forestry,
Syracuse, New York, has accepted a temporary position as expert with the
Office of Forest Pathology, Bureau of Plant Industry. Doctor Penning-
ton will have charge of the season's work on white pine blister rust eradi-
cation in the state of Michigan.
Mr. G. H. Godfrey, of Iowa State College, was appointed scientific
assistant in Cotton, Truck and Forage Crop Disease Investigations,
Bureau of Plant Industry, effective June 8. Mr. Godfrey was formerly
scientific assistant in Cereal Disease Investigations, but during the past
year was granted leave of absence to engage in post-graduate study.
LITERATURE ON PLANT DISEASES
Compiled bt Eunice R. Oberlt, Librarian, Bureau or Pu^nt Induhtrt. akd
Florence P. Smith, Absihtant
April to May, 1917
American Phytoptthological Society. Report of the eighth annual meeting. Phyto-
patholojo' 7: 14,>-149. Ap. 1917.
Pacific Division. Report of meeting .... Phytopathology 7: l.SO-
151. Ap. 1917.
Held at UnivcrHity of California. Rerkelcy, D. 2K and 29. 19!r>.
Anderson, Jacob Peter. Plant diiieases. Alnnka .\gr. Kxpt. Sta. Rpt. 191i: .19 41
1910.
Ashby, S. F. Report on l<*af R|>ot8 of okra and tomato. Jour. Jamaica .\gr Soc.
21: 13. Ja. 1917.
(Vrrospora on okra; Cladonporium fulrum (m tomato.
Australia. Minister of Lands. I)i8ea«efi in plane trees. .Aust. Internat. NurM-rv-
munl6. no. 2: 15. F. 1917.
Ur|M)rt of c'onferenrr relative to arrioufi fungoid diiieaee. which has recently
Imth attacking plane tn*cH.
Avema Sacca, Rosarto. MolcHtiiis cryptogamiriifl do caf6eiro. IM. Agr. i.Sln
Piiulo): 17: S7S 922. ilhw. N. 1910.
MoIcHti:!.** (Typtoganiirafl da canna do aMUcar. R<>1. Agr. (Siko Paulo) 17:
IW-lKiS. illiw. I). 1910.
Bancroft, C. Keith. Tht* Iraf ditvenfip of ruhlMT. CNinditionn in Surinam. Jour.
]\i\. Agr. Hrit. (luiana 10: 93 1(». Ja. 1917.
Beattle, Rollo Kent. TUv intriMlurtion of fon*ign plant diM*a»eii. (.\lwtrairt
Sriinn- n. H. 46: 139. .My. 1. 1917.
Bethel, Ellsworth. Pun-inia Hultnitrnn and ittt iu>fial hunts. I'hytopatholog> 7:
H2 'M. Ap. 1917.
Bols, D^slri. Snnt'>t«' dr |iatltologir v^grtalr. Stance du 2 mars 1917. \U-\-
Unrt 89: "JIM .Mr. 10. 1917.
Brown, Harry Bates. I.ifr Iti^tnry and poisonfiUH pro|»orti«*s of (Mavicciw panpali
Jniir \4!r Ki ..«:irrli 7: 40l t<N>. 2 «« . pi .TJ. N. 27. 1916.
Brunner, Stephen C. Jd'pnrt uf a irip to Pinar did Rio.| .Xgricultun* jCulia' 1:
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Burkholder, Walter H. 'Mh- ]HrfiTt htap* of (ilceoHlNiriuni vi>ni*tiun. Phylopa-
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/*.'»|-.'m»/- ■" f I'/'l \ I 1.1 til II Mp,
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Ki-piirt 4i( thi- lltMMl Kivi-r )ir:iiii-h rxpi-niin lit ittatioh for 1916.
19171 Literature on Plant Diseases 323
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Most common diseases of the year, p. 367-374.
Darnell-Smith, G. P. ''Brown spot" of Emperor mandarins. Elxperiments in
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Ja. 1917.
Literature cited, p. 80.
Bacillus citrimaculans n. sp.
Elliott, John Asbury. The sweet potato "soil rot'' or "pox/' a slime mold disease.
Delaware Agr. Expt. Sta. Bui. 114, 25 p., illus., 5 pi. 1916.
Cystospora batata gen. nov., sp. nov.
Fawcett, Howard S. A bark disease of avocado trees. Ann. Rpt. California Avo-
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Pythiacystis citrophthora.
Freiberg, Geoige W. Studies in the mosaic diseases of plants. Ann. Missouri
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BiWiography, p. 223-225.
Galloway, Beverly Thomas. Newton B. Pierce. Phytopathology 7: 143-144. Ap.
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Died O. 13, 1916 at the age of 60 years.
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Destructive bacterial disease undoubtedly the same as that described in 1897
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Oermano de Souza, Raul. Tylenchus acuto-caudatus nos cafezaes de Indaiatuba.
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Higgins, Bascombe Britt. Notes on some disease of collards. Georgia Agr. Expt.
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Black mold, Altemaria hrasstcce; sclerotinose, Sclerotinia libertiana.
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Decay in timber is almost exclusively due to the action of fungi, the greater
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1917.
Occurrence in United States.
324 Phytopathology [Vol. 7
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Reprinted from Hot. Oaz. Noted in previous list.
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McClintock, James A. Sclerotinia blight, a serioiu* dirM^am* of nnap lN*anji rauM-d
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1917.
Literature, p. 4S5.
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Lite nit lire ritetl. p. 21«V 220.
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Phytop.itliology 7: 101-105. Ap. 1917.
Literature cited, p. 105.
.1/. tnntitirnhi II. .H|».
Matz, J. A RhiKoi tniiia of the fig. I'hytopatliohiKy 7: 110-llH. 3 tig., pi. 2 Ap
11U7.
HibliMKraphif*al fontnoteH.
H. mnrttf^rlrrntm ii. Hp.
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try 15: Jl.VJ.'l I". 1'.»17.
Melsnder, Aiel Leonard, and Heald, Frederick De Forest. 11h* control of fr*itt
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Melhus, IrTing E., and Durrell, L. W. The burlN'rry biif<h and black utmi ni^t uf
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Murphy, Paul A. Ilic iiii'**.ih lii-ia"!* nf |Hitat«H*f». .Xier. (iai. Canada 4: ^i45 .%!*•.
jll'j- M\ l'»17
1917] Literature on Plant Diseases 326
Nowell, William. The fungi of internal boll disease. West Indian Bui. 16: 152-159,
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References, p. 159.
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Dept. of agriculture, Ceylon — Leaflet no. 2.
Hypochnus sp.
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Reddick, Donald. Serious diseases of the season. Proc. West. New York Hort.
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Rosenbaum, Joseph. Studies of the genus Phytophthora. Proc. Nat. Acad. Sci.
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Scott, Charles A. A practical method of preventing the damping off of coniferous
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Shaw, Jacob Kingsley. The varietal relations of crown gall. Science n. s. 46:
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Sherbakoff, Constantin Demetry. Buckeye rot of tomato fruit. Phytopathology
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Phytophthora lerrestria n. sp.
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References in footnotes.
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Stevens, Neil Everett, and Wilcox, R. B. Rhizopus rot of strawberries in transit.
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Literature cited, p. 21-22.
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Physalospora theohrom<B n. sp.
Stachylidium theohromcB n. sp.
Helminthosporium theobromae n. sp.
326 Phytopathology (Vol. 7
U. S. Department of Agriculture. Federml Horticultural Botrd. Service and rrcii-
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Gymnosjtorangium juni j)cri-virginiiitnrt O. glohonum on Juniperum nrgtnft'ii
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PhytopathoIog>' 7: Kio i;n). 2 fig. Ap. 1917.
Bibliographical footnotes.
Recent cultures of f<ire8t tree rUHts. Phytopathology 7: 106 IW
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Stn. Hul. ir»l. 24 p.. illuH. K. 1917.
Hil>liography. p. 22-24.
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Bibliography, p. 243 245.
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Bibliography, p. 154-155.
Running title: Durability of yellow pine.
iPiiYioi'Mii(»LcHiY, for Juiu\ I1M7 '7: \'ut'2'Mi wan iiwued June 2. 1917 ,
PHYTOPATHOLOGY
VOLUME VII NUMBER 5
OCTOBER, 1917
ALTERNARIA ON DATURA AND POTATO
R. D. Rands
With Four Figures in the Text
During the progress of studies on the early blight of potato caused by
AUernaria solani (E. & M.) J. & G., an investigation has been made of the
various host relations attributed to this fungus. Throughout the litera-
ture the early blight organism is claimed to be the cause of the leaf-spot
of Jimson weed {Datura ^ various species). One author (Chester, 1892)
goes so far as to state that this was probably the original host, the fungus
going from it to the potato and tomato. The object of this paper is to
present some comparative studies of early blight and the Datura leaf-spot
in order to clear the apparent misunderstanding of the relationship be-
tween these two diseases.
The Datura leaf-spot has been under the observation of the author in
the University Pharmaceutical Garden, Madison, Wisconsin, during the
summers of 1915 and 1916. The spots show a zonation similar to that
of early blight but they are straw colored rather than deeply stained
(fig. 1). They first appear on the lower more shaded leaves. Under
favorable conditions the disease spreads gradually upwards when finally,
in late autumn, the seed pods often develop dark sunken lesions (fig. 2).
When a leaf becomes weakened by a number of enlarged spots it is usu-
ally shed from the plant.
The disease was noted on the following species and varieties growing in
the Pharmaceutical Garden September 15, 1915:^
Datura tatula Linn., D. tatula inermis, D. stramonium Linn., D. stramon-
ium inermiSj D. stramonium gigardeum, D, inermis Jacq., D, fastuosa Linn.,
D. ferox Linn., D. laeviSj D. querdfolia H. B. K., D. leichardtii F. Muell.,
D. metel Linn.(?).
* The species and varieties are listed in the form in which they were found on the
garden labels. Since the seed was originally of German origin the names are pre-
sumably those in common usage in the foreign seed trade.
328 Phytopathology (Vol. 7
The pod blight was especially conspicuous on D. tatula inermin. ftra-
manium inermis and fastuosa^ while the leaf-spot occurred on all to a
greater or less extent. Datura stramonium giganleum showed grpater
resistance than the others.
ECONOMIC IMPORTANCE OF THE DISEASE
As is well known, the Datune furnish one of the sources of the drug
atropin or daturin which is obtaineil from the leaves. The leaves are
picked in late Septcmlx»r sufficiently early to avoid frost. It is evident
that anything which tends, particularly in the latter part of the 9ew^m.
to reduce the leaf an*a st»riouslv interferes with its economic use. Ki^tiniatrs
made in SeptemlHT 11)15, placnnl the average for all varieties at M) to'iO
per cent of the total leaf surface destroyed. The dr>' season of llUli pn^
ventetl much apiK»araiice of the disease* prior to the rains of SeptenilnT.
But even after that, favorable weather continuing, a loss of 10 to 15 |mt
ct»nt in leaf area occurn'd.
1I18T(»IIY OF TIIK DISEASK AM) ITS CD.NFl'SIOX WITH EARLY HLKiHT
The first refenMic(» inferring that a n^lationship exists l>etwe«»n this*
disease* and early blight is that by (\K)ke ( lS8.*i). He d<»scril)es. as though
they wen* i<ieiitiral. the fungus fn)iii Datura and that fn)m the tomato .v
Macrof<f)ifrium solani (*<M>ke. Me was apimn'iitly unaware that tlu* same
binomial had Im^mi applied by KUis an<l Martin to a simikir fungus ontlie
I)otato the yi'ar U'fon'. Sareanlo (ISStij lists Cooke's fungus as .Wnrri*-
^fHtnutn nn»hi Saec. I>iit4T Kllis (s<»<» .Iom*s ISIW) affirms that this is the
saiiM* fungus whieh he and Martin deseriU'd. Thiw the confu.iion wnmu^
to have originated. It is shown by the follr»wing n'fen»nces which an* a|»-
pan-ntly ronrerned with this Datura leaf-s|K)t. Kellemian (ISHo) fn»!n
Manhattan. Kan>as. Halsted i IS*KV) from New Jer>*ey, Briosi and C'avara
<lV»2i and Terrari** ' P.M^V) fn>m Italv. St4»vens ilSlMi) fnnnOhio. I'mltT-
\mhh\ iin«l liailr lV.»7i and Atkinson ilS*»7) fn>m .\lal)ama, Jon^^ an^i
<Irniit |s*»7i :nnl Orton isiHji funi V«Tniont. So far as the hteraiure
n'\i:il> thr ab«»vr determinations were bas4'd entin*ly on liiorphol«»|cir:d
evitli-rni and on tin- Niinilarity of the <lis4'ast» to early blight of pi»tat«»
an<l iiMt nil iii<>iiil:itinn work with the causal organism.
In onl« r to ilrtiTniinr wliether the leaf->|><tt as found at Madinm i*^
iIh- ^aiiif a* that r('|Nirt4'd i-UrwIuTe. ex>i«*eat:M' lalti'lkHl .W#ifriv/h»»-;:*Fn
.^"'' i . (V M. on Datura. Were e\annnt>d and eoni|KinMl with typir:tl
in;it«i.:il lol'ii . i«i| JM-rr. Tlif ni.irroMopie apfN-araiiee of the S|X>ts of tlif»
<\ .'.i^ii- ni Si\nHi;ir ami I-irlr No. lilo, and other eollection?» fn»m
1917] Rands: Alternaria on Datura and Potato 331
atomized with a spore suspension in water and for forty-eight hours kept
moist by a fine spray from a nozzle. In most eases reisolations made
from the infected plants were successful.
On September 4, 1916, further inoculations with spores of AUernaria
solani were made on mature potato and Datura plants growing in the
field. Several leaves were atomized with a heavy spore suspension, while
with others spores were introduced into pimctures. Successful reisola-
tions of the fungus was secured from both atomized and needle punctured
leaves on the potato. In one instance tissue plantings from the browned
needle punctures on Datura stramonium five weeks after inoculation gave
the fungus. The results corroborate the green house tests and show that,
in no case, was A. solani able to form spots on even the old and weakened
leaves of Jimson weed. However in toto fixations showed that penetration
and incipient infection occurred in many .cases. But the fungus seemed
unable to establish itself and bring about enlargement of the spot.
inoculations with the fungus from datura
The Madison culture of the parasite from Datura stramonium was tested
comparatively first in the greenhouse, and later in the field on Jimson
weed, potato, tomato, and on Solanum nigrum, the common black night-,
shade. Several methods of inoculation in which myceUum or spores wei'e
placed in needle pricks and the spores atomized upon the surface were
used. Most of the tests were carried out at the same time and under the
same conditions as those already reported with Alternaria solani from
potato. These experiments may be briefly summarized as follows: On
the Jimson weeds including Datura stramonium, D. inermis, and D. tatula,
typical spots 3 to 10 mm. in diameter invariably resulted after two weeks.
On potato, tomato and Solanum nigrum, incipient infections in the form
of tiny brown specks often occurred. They were less abundant on the
nightshade and vigorous leaves of tomato. These spots in no case en-
larged, even after the leaves were yellowing and dying. The fungus was
reisolated readily from the Datura leaves but only in a few cases could it
be obtained from the incipient spots on the other plants. The needle
punctures on the latter were in most cases entirely healed after eight days.
Therefore, it appears that we have here an Alternaria which, though bear-
ing much superficial resemblance to AUernaria solani, is nevertheless dis-
tinct in its host relationship. In no instance has there been observed any
crossing over to the potato or other hosts of A, solani tested, and also no
crossing of the potato fungus to Jimson weed. The above conclusion is
further confirmed by a comparison of the two fungi in other particulars.
i
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Rands: Aia^bnaria on DATxmA. and Potato
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334 Phytopathology (Vol. 7
comparison of the fl^gus from datura with alternaria 80l.\m
On hard potato agar the colony of Altcrnaria from Datura is at first
of a light olivacious color with faint pinkish margins; later the entire
growth becomes a grayish white while from the lower surface often blackish
mycelium can be seen through the agar. With A. 9olani the grayi^h or
brownish colony produces a deep pinkish to yellowit*h pigmentation of the
agar which often extends in advance of the mycelium (fig. 3). This fur-
nishes at once a distinct physiological ba.^is for the differentiation of the
two fungi. Morphologically they are similar in many respects. <>ne
of the most prominent characters which distinguishes the Datura f unguis
from A, i<olani is found in the terminal prolongation or beak of the spore.
In the former the beak is coarser, more elongated, rarely has any pro-
nounced tapering to the tip, and is never forked or divided (fig. 4). ( >ften
a.** high as 75 per cent of the spores of -4. solmii l)oth from cultures and
from s|)ots, have l)eeii ol)ser\'ed to possess forked or variously di\'i*kHl
l)eaks, the sulnlivisions ta|Kring more or less to the tip. One hundred
HIK)res from typical early blight s|X)ts from potato gave a range in Mie erf
12() to 2<U) by 12 to 20 /i and an average of 2(K) by 17 fi. With the Datura
fungus the range in size was 128 to 448 by It) to 40 m with an average of
2t)l by 23 /i- 'i'he latter it is sc*<*n has spores considerably larger in both
dimensions than -4. W/iri/. However, the difference is not such as to
make sjwre measureineiits n»liable when not supplemented by the oth«*r
distinguishing characters mentione<l.
IDENTITY OK THE DATURA PARASITE
The fungus has U»imi refern»<l to as an Altemaria. This was. howeviT.
inen*ly on account of its great similarity to Aliernaria nolani. As in the
(*as4' of the latter, cultun^s. on oatnieal agar oi*casionally develop f(port> in
catcnulute pairs (fig. 4). Acconling to the pn^sent delimitations of ttie
g(*n<Ta Alt4*nmria and Macros|)oriuni this catenulation of spores make:« the
fungus an Alt4*rnaria. However, it is n^aliauMl that the conditioa< under
wlkirh s])on' i>airs are fonned an* |M>ssibly abnonnal and it is doubtful
if the